Search Results (152)

In this study, the functions of the photosynthetic machinery were evaluated using chlorophyll a fluorescence technique (PAM and JIP test) in pea plants (Pisum sativum L. cv Borec) and its LHC II oligomerization variants (mutants Costata 2/133 and Coeruleovireus 2/16). The oligomeric forms of LHCII increased in the following order: Costata 2/133 < Borec wt < Coeruleovireus 2/16. Data revealed that the mutant with higher LHCII oligomerization (Coeruleovireus 2/16) at low light intensity (LL, 150 µmol photons/m²·s) exhibited the following: (i) decreased energy dissipation and increased electron transport efficiency; (ii) higher reaction center density; (iii) increased amounts of the open reaction centers (qp) and their excitation efficiency (Φexc); and (iv) influenced the reoxidation of Q_A⁻, alleviating its interaction with plastoquinone. These effects enhanced photosynthetic performance related to PSII photochemistry (PI_ABS) and overall photosynthetic efficiency (PItotal). High light intensity (HL, 500 µmol photons/m²·s) caused a reduction in open reaction centers (qp), excitation efficiency (Φexc), photochemical energy conversion of PSII (Φ_PSII), maximum efficiency of PSII photochemistry in light (Fv′/Fm′), and linear electron transport via PSII, with more pronounced effects observed in membranes with a lower degree of LHCII oligomerization (Costata 2/133). This study provides novel experimental evidence for the pivotal role of the LHCII structural organization in determining the efficiency of light-dependent reactions of photosynthesis. Full article

The fraction of open Photosystem II (PSII) reaction centers (q_L) is critical for connecting broadband PSII fluorescence (ChlF_PSII) with the actual electron transport from PSII to Photosystem I. Accurately estimating q_L is fundamental for determining ChlF_PSII, which, in turn, is vital for mechanistically estimating the actual electron transport rate and photosynthetic CO₂ assimilation. Chlorophyll fluorescence provides direct physiological insights, offering a robust foundation for q_L estimation. However, uncertainties in the ChlF_PSII–q_L relationship across different plant functional types (PFTs) limit its broader application at large spatial scales. To address this issue, we developed a leaf-level instrument capable of simultaneously measuring actively and passively induced chlorophyll fluorescence. Using this system, we measured light response, CO₂ response, and temperature response curves across 52 species representing seven PFTs. Our findings reveal the following: (1) a strong linear correlation between ChlF_PSII derived from passively induced fluorescence and that from actively induced fluorescence (R² = 0.85), and (2) while the parameters of the ChlF_PSII–q_L relationship varied among PFTs, ChlF_PSII reliably modeled q_L within each PFT, with the R² ranging from 0.85 to 0.96. This study establishes quantitative ChlF_PSII–q_L relationships for various PFTs by utilizing passively induced fluorescence to calculate ChlF_PSII. The results demonstrate the potential for remotely sensed chlorophyll fluorescence data to estimate q_L and strengthen the use of fluorescence-based approaches for mechanistic GPP estimation at large spatial scales. Full article

The tomato is among the crops with the most extensive cultivated area and greatest consumption in our nation; nonetheless, secondary salinization of facility soil significantly hinders the sustainable growth of facility agriculture. Melatonin (MT), as an innovative plant growth regulator, is essential in stress responses. This research used a hydroponic setup to replicate saline stress conditions. Different endogenous levels of melatonin (MT) were established by foliar spraying of 100 μmol·L⁻¹ MT, the MT synthesis inhibitor p-CPA (100 μmol·L⁻¹), and a combination of p-CPA and MT, to investigate the mechanism by which MT mitigates the effects of salt stress on the photosynthetic efficiency of tomato seedlings. Results indicated that after six days of salt stress, the endogenous MT content in tomato seedlings drastically decreased, with declines in the net photosynthetic rate and photosystem performance indices (PI_total and PI_abs). The OJIP fluorescence curve exhibited distortion, characterized by anomalous K-band and L-band manifestations. Exogenous MT dramatically enhanced the gene (TrpDC, T5H, SNAcT, and AcSNMT) expression of critical enzymes in MT synthesis, therefore boosting the level of endogenous MT. The application of MT enhanced the photosynthetic parameters. MT treatment decreased the fluorescence intensities of the J-phase and I-phase in the OJIP curve under salt stress, attenuated the irregularities in the K-band and L-band performance, and concurrently enhanced quantum yield and energy partitioning ratios. It specifically elevated φP_o, φE_o, and ψ_o, while decreasing φD_o. The therapy enhanced parameters of both the membrane model (ABS/RC, DI_o/RC, ET_o/RC, and TR_o/RC) and leaf model (ABS/CS_m, TR_o/CS_m, ET_o/CS_m, and DI_o/CS_m). Conversely, the injection of exogenous p-CPA exacerbated salt stress-related damage to the photosystem of tomato seedlings and diminished the beneficial effects of MT. The findings suggest that exogenous MT mitigates salt stress-induced photoinhibition by (1) modulating endogenous MT concentrations, (2) augmenting PSII reaction center functionality, (3) safeguarding the oxygen-evolving complex (OEC), (4) reinstating PSI redox potential, (5) facilitating photosynthetic electron transport, and (6) optimizing energy absorption and dissipation. As a result, MT markedly enhanced photochemical performance and facilitated development and salt stress resilience in tomato seedlings. Full article

Aspirin (Asp) is extensively used in human health as an anti-inflammatory, antipyretic, and anti-thrombotic drug. In this study, we investigated if the foliar application of Asp on tomato plants has comparable beneficial effects on photosynthetic function to that of salicylic acid (SA), with which it shares similar physiological characteristics. We assessed the consequences of foliar Asp-spray on the photosystem II (PSII) efficiency of tomato plants, and we estimated the reactive oxygen species (ROS) generation and the chloroplast ultrastructural changes. Asp acted as an osmoregulator by increasing tomato leaf water content and offering antioxidant protection. This protection kept the redox state of plastoquinone (PQ) pull (qp) more oxidized, increasing the fraction of open PSII reaction centers and enhancing PSII photochemistry (Φ_PSII). In addition, Asp foliar spray decreased reactive oxygen species (ROS) formation, decreasing the excess excitation energy on PSII. This resulted in a lower singlet oxygen (¹O₂) generation and a lower quantum yield for heat dissipation (Φ_NPQ), indicating the photoprotective effect provided by Asp, especially under excess light illumination. Simultaneously, we observed a decrease in stomatal opening by Asp, which reduced the transpiration. Chloroplast ultrastructural data revealed that Asp, by offering a photoprotective effect, decreased the need for the photorespiration process, which reduces photosynthetic performance. It is concluded that Asp shares similar physiological characteristics with SA, having an equivalent beneficial impact to SA by acting as a biostimulant of the photosynthetic function for an enhanced crop yield. Full article

The success of establishing new trees in cities and their subsequent growth depend, among others, on the proper selection of tree species which can easily tolerate the post-planting stress. In the spring of 2023, young Italian alder (Alnus cordata (Loisel.) Duby) and common lime (Tilia × europaea L. ‘Pallida’) trees were planted in a street of heavy traffic in Warsaw. In the summer of 2023, leaf samples were collected during the growing season for chlorophyll a fluorescence measurements and chemical analyses. Additionally, the autumn phenological phases were monitored. Chlorophyll a fluorescence measurements revealed higher values of F_v/F_m, density of reaction centers per cross-section, and electron transport chain efficiency between photosystems II and I, as well as lower energy dissipation rate per active reaction center of photosystem II in A. cordata. Moreover, A. cordata revealed higher chlorophyll a, chlorophyll b, and carotenoid content. The flavonoid and proline content in both species was the highest by the end of July and then decreased. In T. × europea ‘Pallida’, the contents of these stress biomarkers increased in the late growing season. Our results showed that T. × europaea ‘Pallida’ is less resistant to post-planting stress in urban conditions, while A. cordata showed higher resistance to variable weather conditions, high photosynthetic efficiency, and long foliage lifespan. Full article

Dehydration is known to affect the rate of electron transfer backreaction from the light-induced charge separation state P⁺Q_A⁻ to the neutral ground state PQ_A in photosynthetic bacterial Reaction Centers. On the other hand, a 20 s continuous illumination period has been demonstrated to induce (at 297 K) formation of one or more light-adapted states at different levels of dehydration; these light-adapted states are believed to be related to peculiar response(s) from the protein. In this work, we applied time-resolved rapid-scan FTIR difference spectroscopy to investigate the protein response under dehydrated conditions (RH = 11%) at 281 K both after a flash and under prolonged continuous illumination. Time-resolved FTIR difference spectra recorded after a laser flash show a protein recovery almost synchronous to the electron transfer backreaction P⁺Q_A⁻ → PQ_A. Time-resolved FTIR difference spectra recorded after 20.5 s of continuous illumination (RH = 11%, T = 281 K) surprisingly show almost the same kinetics of electron transfer back reaction compared to spectra recorded after a laser flash. This means that the mechanism of formation of a light-adapted stabilized state is less effective compared to the same hydration level at 297 K and to the RH = 76% hydration level (both at 281 K and 297 K). Time-resolved FTIR difference spectra after continuous illumination also suggest that the 1666 cm⁻¹ protein backbone band decays faster than marker bands for the electron transfer back reaction P⁺Q_A⁻ → PQ_A. Finally, FTIR double-difference spectra (FTIR difference spectrum recorded after 18.4 s illumination minus flash-induced FTIR difference spectrum) suggest that at RH = 11%, a light-adapted state different from the one observed at RH = 76% is formed. A possible interpretation is that at RH = 11%, the protein response is modified by the fact that only protons can move easily, differently from water molecules, as instead observed for RH = 76%. This probably makes the formation of a real light-adapted P⁺Q_A⁻ stabilized state at RH = 11% unfeasible. Full article

1-Methyl cyclopropene (1-MCP) is known as an ethylene antagonist, yet its mechanisms in regulating photosynthetic electron transport and energy dissipation in chrysanthemum under heat stress are not well understood. Here, the chlorophyll a fluorescence and modulated 820 nm reflection transients were analyzed in heat-tolerant and heat-sensitive chrysanthemum plants. This study demonstrates that 1-MCP pre-treatment helps maintain the net photosynthetic rate (P_n) and the reaction center activity of photosystems I and II (PSI and PSII) during heat stress. Specifically, 1-MCP treatment significantly increases the fraction of active oxygen-evolving complex (OEC) centers and reduces relative variable fluorescence intensity at the J step (V_J) as well as the efficiency of electron transfer at the PSI acceptor side (δ_Ro). These effects mitigate damage to the photosynthetic electron transport chain. Additionally, 1-MCP-treated plants exhibit decreased quantum yield of energy dissipation (φ_Do) and reduced energy flux per reaction center (DI_o/RC). Overall, 1-MCP enhances light utilization efficiency and excitation energy dissipation in the PSII antennae, alleviating heat stress-induced damage to PSI and PSII structures and functions. This study not only advances our understanding of 1-MCP’s regulatory role in photosynthetic processes under heat stress but also provides a basis for using exogenous substances to improve chrysanthemum heat resistance. Full article

The relative impacts of biochemical and stomatal limitations on photosynthesis during photosynthetic induction have been well studied for diverse plants under ambient CO₂ concentration (C_a). However, a knowledge gap remains regarding how the various photosynthetic components limit duction efficiency under elevated CO₂. In this study, we experimentally investigated the influence of elevated CO₂ (from 400 to 800 μmol mol^–1) on photosynthetic induction dynamics and its associated limitation components in two broadleaved tree species, Populus tomentosa and Eucalyptus robusta. The results show that elevated CO₂ increased the steady-state photosynthesis rate (A) and decreased stomatal conductance (g_s) and the maximum carboxylation rate (V_cmax) in both species. While E. robusta exhibited a decrease in the linear electron transport rate (J) and the fraction of open reaction centers in photosynthesis II (q_L), P. tomentosa showed a significant increase in non-photochemical quenching (NPQ). With respect to non-steady-state photosynthesis, elevated CO₂ significantly reduced the induction time of A following a shift from low to high light intensity in both species. Time-integrated limitation analysis during induction revealed that elevated CO₂ reduces the relative impacts of stomatal limitations in both species, consequently shifting the predominant limitation on induction efficiency from stomatal to biochemical components. Additionally, species-specific changes in q_L and NPQ suggest that elevated CO₂ may increase biochemical limitation by affecting energy allocation between carbon fixation and photoprotection. These findings suggest that, in a future CO₂-rich atmosphere, plants productivity under fluctuating light may be primarily constrained by photochemical and non-photochemical quenching. Full article

To sustain agricultural productivity and safeguard global food security, and confront the escalating challenges posed by climate change and water scarcity, it is essential to enhance the growth and productivity of rice under water stress. This study investigated the effects of lanthanum chloride on the chlorophyll fluorescence characteristics and grain yield of rice under different irrigation modes. The rice cultivar H You 518 was selected and sprayed 20, 100, or 200 mg·L⁻¹ lanthanum chloride at the booting and heading stages under deficit irrigation (where no rewatering was applied after the initiation of stress, allowing the water layer to evaporate naturally under high temperatures) or conventional irrigation (with daily rewatering to maintain a consistent water level). The results showed that the application of low concentrations lanthanum chloride promoted the chlorophyll content, whereas high concentrations decreased the chlorophyll content, under deficit irrigation, the effect of lanthanum chloride on the green fluorescence parameters of rice leaves at the booting stage was greater than that at the heading stage, and the booting stage was more sensitive to water deficit. The application of 100 mg·L⁻¹ lanthanum chloride reduced the initial fluorescence (F0) and the non-photochemical quenching coefficient (qN); promoted the activity of leaf photosynthetic system II (PSII); and maximized the photochemical quantum yield (Fv/Fm), photochemical quenching coefficient (qP), and PSII relative electron transfer efficiency (ETR). Under deficit irrigation, this treatment significantly enhanced grain yield by increasing the thousand-grain weight, spikelet filling rate, and number of grains per panicle. These results suggest that spraying 100 mg·L⁻¹ lanthanum chloride at the booting stage under deficit irrigation can effectively increase the chlorophyll content, thereby increasing the light energy conversion efficiency of the PS II reaction center, ultimately resulting in increased spikelet filling rate and grain yields. Full article

Disaccharide trehalose has been proven in many cases to be particularly effective in preserving the functional and structural integrity of biological macromolecules. In this work, we studied its effect on the electron transfer reactions that occur in the chromatophores of the photosynthetic bacterium Cereibacter sphaeroides. In the presence of a high concentration of trehalose, following the activation of the photochemistry by flashes of light, a slowdown of the electrogenic reactions related to the activity of the photosynthetic reaction center and cytochtome (cyt) bc₁ complexes is observable. The kinetics of the third phase of the electrochromic carotenoid shift, due to electrogenic events linked to the reduction in cyt b_H heme via the low-potential branch of the cyt bc₁ complex and its oxidation by quinone molecule on the Q_i site, is about four times slower in the presence of trehalose. In parallel, the reduction in oxidized cyt (c₁ + c₂) and high-potential cyt b_H are strongly slowed down, suggesting that the disaccharide interferes with the electron transfer reactions of the high-potential branch of the bc₁ complex. A slowing effect of trehalose on the kinetics of the electrogenic protonation of the secondary quinone acceptor Q_B in the reaction center complex, measured by direct electrometrical methods, was also found, but was much less pronounced. The direct detection of carbohydrate content indicates that trehalose, at high concentrations, permeates the membrane of chromatophores. The possible mechanisms underlying the observed effect of trehalose on the electron/proton transfer process are discussed in terms of trehalose’s propensity to form strong hydrogen bonds with its surroundings. Full article

The fundamental key to increase photosynthetic efficiency of crop plants lies in optimizing the light energy use efficiency. In our study, we used tomato to evaluate the allocation of absorbed light energy in young and mature leaves, and to estimate if the extent of photoinhibition and photoprotection can be affected by the leaf age. A reduced efficiency of the oxygen-evolving complex, in young leaves compared to mature ones, resulted in a donor-side photoinhibition, as judged from the significantly lower Fv/Fm ratio, in young leaves. The detected increased ¹O₂ production in young leaves was probably due to a donor-side photoinhibition. The effective quantum yield of photosystem II (PSII) photochemistry (Φ_PSII), at low light intensity (LLI, 426 μmol photons m⁻² s⁻¹), was significantly lower in young compared to mature leaves. Moreover, the non-significant increase in non-photochemical energy loss in PSII (Φ_NPQ) could not counteract the decreased Φ_PSII, and as a result the non-regulated energy loss in PSII (Φ_NO) increased in young leaves, compared to mature ones. The significantly lower Φ_PSII in young leaves can be attributed to the increased reactive oxygen species (ROS) creation that diminished the efficiency of the open PSII reaction centers (Fv’/Fm’), but without having any impact on the fraction of the open reaction centers. The reduced excess excitation energy, in mature leaves compared to young ones, at LLI, also revealed an enhanced PSII efficiency of mature leaves. However, there was almost no difference in the light energy use efficiency between young and mature leaves at the high light intensity (HLI, 1000 μmol photons m⁻² s⁻¹). The ability of mature tomato leaves to constrain photoinhibition is possible related to an enhanced photosynthetic function and a better growth rate. We concluded that the light energy use efficiency in tomato leaves is influenced by both the leaf age and the light intensity. Furthermore, the degrees of photoinhibition and photoprotection are related to the leaf developmental stage. Full article

The pH dependence of the free energy level of the flash-induced primary charge pair P⁺I_A⁻ was determined by a combination of the results from the indirect charge recombination of P⁺Q_A⁻ and from the delayed fluorescence of the excited dimer (P*) in the reaction center of the photosynthetic bacterium Rhodobacter sphaeroides, where the native ubiquinone at the primary quinone binding site Q_A was replaced by low-potential anthraquinone (AQ) derivatives. The following observations were made: (1) The free energy state of P⁺I_A⁻ was pH independent below pH 10 (–370 ± 10 meV relative to that of the excited dimer P*) and showed a remarkable decrease (about 20 meV/pH unit) above pH 10. A part of the dielectric relaxation of the P⁺I_A⁻ charge pair that is not insignificant (about 120 meV) should come from protonation-related changes. (2) The single exponential decay character of the kinetics proves that the protonated/unprotonated P⁺I_A⁻ and P⁺Q_A⁻ states are in equilibria and the rate constants of protonation k_on^H +k_off^H are much larger than those of the charge back reaction k_back ~10³ s⁻¹. (3) Highly similar pH profiles were measured to determine the free energy states of P⁺Q_A⁻ and P⁺I_A⁻, indicating that the same acidic cluster at around Q_B should respond to both anionic species. This was supported by model calculations based on anticooperative proton distribution in the cluster with key residues of GluL212, AspL213, AspM17, and GluH173, and the effect of the polarization of the aqueous phase on electrostatic interactions. The larger distance of I_A⁻ from the cluster (25.2 Å) compared to that of Q_A⁻ (14.5 Å) is compensated by a smaller effective dielectric constant (6.5 ± 0.5 and 10.0 ± 0.5, respectively). (4) The P* → P⁺Q_A⁻ and I_A⁻Q_A → I_AQ_A⁻ electron transfers are enthalpy-driven reactions with the exemption of very large (>60%) or negligible entropic contributions in cases of substitution by 2,3-dimethyl-AQ or 1-chloro-AQ, respectively. The possible structural consequences are discussed. Full article

In addition to leaves, photosynthesis can occur in other green plant organs, including developing seeds of many crops. While the majority of studies examining photosynthesis are concentrated on the leaf level, the role of other green tissues in the production of total photoassimilates has been largely overlooked. The present work studies the photosynthetic behavior of leaves and non-foliar (pericarps, coats, and cotyledons) organs of pea (Pisum sativum L.) plants at the middle stage of seed maturation. The Chl a fluorescence transient was examined based on OJIP kinetics using the FluorPen FP 110. A discrepancy was observed between the performance index (PI_ABS) for foliar and non-foliar plant tissues, with the highest level noted in the leaves. The number of absorbed photons (ABS) and captured energy flow (TRo) per reaction center (RC) were elevated in the non-foliar tissues, which resulted in a faster reduction in Q_A. Conversely, the energy dissipation flux per RC (DIo/RC and PHI_Do) indicated an increase in the overall dissipation potential of active reaction centers of photosystem II. This phenomenon was attributed to the presence of a higher number of inactive RCs in tissues that had developed under low light intensity. Furthermore, the expression of genes associated with proteins and enzymes that regulate ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBisCo) activity was observed, including chaperonins Cpn60α and Cpn60β, RuBisCO activase, as well as phosphoribulokinase. The expression of these genes was found to differ between foliar and non-foliar tissues, indicating that the activation state of RuBisCO may be modified in response to light intensity. Overall, the present study provides insights into the mechanisms by which non-foliar green tissues of plants adapt to efficient light capture and utilization under low light conditions. Full article

Breeding high-photosynthetic-efficiency wheat varieties is a crucial link in safeguarding national food security. Traditional identification methods necessitate laborious on-site observation and measurement, consuming time and effort. Leveraging unmanned aerial vehicle (UAV) remote sensing technology to forecast photosynthetic indices opens up the potential for swiftly discerning high-photosynthetic-efficiency wheat varieties. The objective of this research is to develop a multi-stage predictive model encompassing nine photosynthetic indicators at the field scale for wheat breeding. These indices include soil and plant analyzer development (SPAD), leaf area index (LAI), net photosynthetic rate (Pn), transpiration rate (Tr), intercellular CO₂ concentration (Ci), stomatal conductance (Gsw), photochemical quantum efficiency (PhiPS2), PSII reaction center excitation energy capture efficiency (Fv’/Fm’), and photochemical quenching coefficient (qP). The ultimate goal is to differentiate high-photosynthetic-efficiency wheat varieties through model-based predictions. This research gathered red, green, and blue spectrum (RGB) and multispectral (MS) images of eleven wheat varieties at the stages of jointing, heading, flowering, and filling. Vegetation indices (VIs) and texture features (TFs) were extracted as input variables. Three machine learning regression models (Support Vector Machine Regression (SVR), Random Forest (RF), and BP Neural Network (BPNN)) were employed to construct predictive models for nine photosynthetic indices across multiple growth stages. Furthermore, the research conducted principal component analysis (PCA) and membership function analysis on the predicted values of the optimal models for each indicator, established a comprehensive evaluation index for high photosynthetic efficiency, and employed cluster analysis to screen the test materials. The cluster analysis categorized the eleven varieties into three groups, with SH06144 and Yannong 188 demonstrating higher photosynthetic efficiency. The moderately efficient group comprises Liangxing 19, SH05604, SH06085, Chaomai 777, SH05292, Jimai 22, and Guigu 820, totaling seven varieties. Xinmai 916 and Jinong 114 fall into the category of lower photosynthetic efficiency, aligning closely with the results of the clustering analysis based on actual measurements. The findings suggest that employing UAV-based multi-source remote sensing technology to identify wheat varieties with high photosynthetic efficiency is feasible. The study results provide a theoretical basis for winter wheat phenotypic monitoring at the breeding field scale using UAV-based multi-source remote sensing, offering valuable insights for the advancement of smart breeding practices for high-photosynthetic-efficiency wheat varieties. Full article

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 (Φ_PSII), 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. Full article