Zinc (Zn) is an essential trace element but can lead to water contamination and ecological deterioration when present in excessive amounts. Therefore, investigating the photosynthetic response of microalgae to Zn stress is of great significance. In this study, we assessed the photosynthetic responses of neutrophilic Chlamydomonas reinhardtii and acidophilic Chlamydomonas sp. 1710 to Zn exposure for 96 h. The specific growth rate (μ), chlorophyll-a (Chl-a) content, and chlorophyll fluorescence parameters were determined. The results demonstrated that Chlamydomonas sp. 1710 was much more tolerant to Zn than C. reinhardtii, with the half-maximal inhibitory concentration (IC50) values of 225.4 mg/L and 23.4  mg/L, respectively. The μ and Chl-a content of C. reinhardtii decreased in the presence of 15  mg/L Zn, whereas those of Chlamydomonas sp. 1710 were unaffected by as high as 100  mg/L Zn. Chlorophyll fluorescence parameters indicated that the regulation of energy dissipation, including non-photochemical quenching, played a crucial role in Zn stress resistance for both Chlamydomonas strains. However, in the case of C. reinhardtii, non-photochemical quenching was inhibited by 5  mg/L Zn in the first 48 h, whereas for Chlamydomonas sp. 1710, it remained unaffected under 100  mg/L Zn. Chlamydomonas sp. 1710 also exhibited a 20 times stronger capacity for regulating the electron transfer rate than C. reinhardtii under Zn stress. The light energy utilization efficiency (α) of Chlamydomonas sp. 1710 had the most highly non-linear correlation with μ, indicating the energy utilization and regulation process of Chlamydomonas sp. 1710 was well protected under Zn stress. Collectively, our findings demonstrate that the photosystem of Chlamydomonas sp. 1710 is much more resilient and tolerant than that of C. reinhardtii under Zn stress.

Lichens are poikilohydric organisms and an important part of the ecosystem. They show high desiccation tolerance, but the mechanism of dehydration resistance still needs to be studied. The photosynthesis recovery of the photobiont in rehydrated lichen Cladonia stellaris after 11-year desiccation was investigated by simultaneously monitoring both photosystem I and II (PSI and PSII) activities. The responses of the photochemical efficiency and relative electron transport rate (rETR) of PSI and PSII, and the quantum yield of the cyclic electron flow (CEF) were measured using a Dual-PAM-100 system. PSI recovered rapidly, but PSII hardly recovered in C. stellaris during rehydration. The maximal photochemical efficiency of PSII (Fv/Fm) was generally very low and reached about just 0.4 during the rehydration. These results indicated that PSII had restored little and was largely inactivated during rehydration. The quantum yield of PSI recovered quickly to almost 0.9 within 4 h and remained constant at nearly 1 thereafter. The results showed that the activation of the CEF in the early stages of rehydration helped the rapid recovery of PSI. The quantum yield of the CEF made up a considerable fraction of the quantum yield of PSI during rehydration. A regulated excess energy dissipation mechanism and non-photochemical quenching (NPQ) also recovered. However, the small extent of the recovery of the NPQ was not enough to dissipate the excess energy during rehydration, which may be responsible for the weak activity of PSII during rehydration. The results indicated that both CEF and NPQ were essential during the rehydration of the photobiont in C. stellaris. The methods used in the measurements of chlorophyll a fluorescence and P700+ absorbance changes in this study provided a speedy and simple way to detect the physiological characteristics of the photobionts of lichen during rehydration. This work improves our understanding of the mechanism behind lichen\'s desiccation tolerance.

Plants often need to withstand multiple types of environmental stresses (e.g., salt and low temperature stress) because of their sessile nature. Although the physiological responses of plants to single stressor have been well-characterized, few studies have evaluated the extent to which pretreatment with non-lethal stressors can maintain the photosynthetic performance of plants in adverse environments (i.e., acclimation-induced cross-tolerance). Here, we studied the effects of sodium chloride (NaCl) pretreatment on the photosynthetic performance of tomato plants exposed to low temperature stress by measuring photosynthetic and chlorophyll fluorescence parameters, stomatal aperture, chloroplast quality, and the expression of stress signaling pathway-related genes. NaCl pretreatment significantly reduced the carbon dioxide assimilation rate, transpiration rate, and stomatal aperture of tomato leaves, but these physiological acclimations could mitigate the adverse effects of subsequent low temperatures compared with non-pretreated tomato plants. The content of photosynthetic pigments decreased and the ultra-microstructure of chloroplasts was damaged under low temperature stress, and the magnitude of these adverse effects was alleviated by NaCl pretreatment. The quantum yield of photosystem I (PSI) and photosystem II (PSII), the quantum yield of regulatory energy dissipation, and non-photochemical energy dissipation owing to donor-side limitation decreased following NaCl treatment; however, the opposite patterns were observed when NaCl-pretreated plants were exposed to low temperature stress. Similar results were obtained for the electron transfer rate of PSI, the electron transfer rate of PSII, and the estimated cyclic electron flow value (CEF). The production of reactive oxygen species induced by low temperature stress was also significantly alleviated by NaCl pretreatment. The expression of ion channel and tubulin-related genes affecting stomatal aperture, chlorophyll synthesis genes, antioxidant enzyme-related genes, and abscisic acid (ABA) and low temperature signaling-related genes was up-regulated in NaCl-pretreated plants under low temperature stress. Our findings indicated that CEF-mediated photoprotection, stomatal movement, the maintenance of chloroplast quality, and ABA and low temperature signaling pathways all play key roles in maintaining the photosynthetic capacity of NaCl-treated tomato plants under low temperature stress.

Photosynthesis is the largest mass- and energy-conversion process on Earth, and it is the material basis for almost all biological activities. The efficiency of converting absorbed light energy into energy substances during photosynthesis is very low compared to theoretical values. Based on the importance of photosynthesis, this article summarizes the latest progress in improving photosynthesis efficiency from various perspectives. The main way to improve photosynthetic efficiency is to optimize the light reactions, including increasing light absorption and conversion, accelerating the recovery of non-photochemical quenching, modifying enzymes in the Calvin cycle, introducing carbon concentration mechanisms into C3 plants, rebuilding the photorespiration pathway, de novo synthesis, and changing stomatal conductance. These developments indicate that there is significant room for improvement in photosynthesis, providing support for improving crop yields and mitigating changes in climate conditions.

Orange carotenoid protein (OCP) is a photoactive protein that participates in the photoprotection of cyanobacteria. There are 2 full-length OCP proteins, 4 N-terminal paralogs (helical carotenoid protein [HCP]), and 1 C-terminal domain-like carotenoid protein (CCP) found in Nostoc flagelliforme, a desert cyanobacterium. All HCPs (HCP1 to 3 and HCP6) from N. flagelliforme demonstrated their excellent singlet oxygen quenching activities, in which HCP2 was the strongest singlet oxygen quencher compared with others. Two OCPs, OCPx1 and OCPx2, were not involved in singlet oxygen scavenging; instead, they functioned as phycobilisome fluorescence quenchers. The fast-acting OCPx1 showed more effective photoactivation and stronger phycobilisome fluorescence quenching compared with OCPx2, which behaved differently from all reported OCP paralogs. The resolved crystal structure and mutant analysis revealed that Trp111 and Met125 play essential roles in OCPx2, which is dominant and long acting. The resolved crystal structure of OCPx2 is maintained in a monomer state and showed more flexible regulation in energy quenching activities compared with the packed oligomer of OCPx1. The recombinant apo-CCP obtained the carotenoid pigment from holo-HCPs and holo-OCPx1 of N. flagelliforme. No such carotenoid transferring processes were observed between apo-CCP and holo-OCPx2. The close phylogenetic relationship of OCP paralogs from subaerial Nostoc species indicates an adaptive evolution toward development of photoprotection: protecting cellular metabolism against singlet oxygen damage using HCPs and against excess energy captured by active phycobilisomes using 2 different working modes of OCPx.

Bundle sheath cells play a crucial role in photosynthesis in C4 plants, but the structure and function of photosystem II (PSII) in these cells is still controversial. Photoprotective roles of bundle sheath chloroplasts at the occurrence of environmental stresses have not been investigated so far. Non-photochemical quenching (NPQ) of chlorophyll a fluorescence is the photoprotective mechanism that responds to a changing energy balance in chloroplasts. In the present study, we found a much higher NPQ in bundle sheath chloroplasts than in mesophyll chloroplasts under a drought stress. This change was accompanied by a more rapid dephosphorylation of light-harvesting complex II (LHCII) subunits and a greater increase in PSII subunit S (PsbS) protein abundance than in mesophyll cell chloroplasts. Histochemical staining of reactive oxygen species (ROS) suggested that the high NPQ may be one of the main reasons for the lower accumulation of ROS in bundle sheath chloroplasts. This may maintain the stable functioning of bundle sheath cells under drought condition. These results indicate that the superior capacity for dissipation of excitation energy in bundle sheath chloroplasts may be an environmental adaptation unique to C4 plants.

Light intensity is highly heterogeneous in nature, and plants have evolved a series of strategies to acclimate to dynamic light due to their immobile lifestyles. However, it is still unknown whether there are differences in photoprotective mechanisms among different light-demanding plants in response to dynamic light, and thus the role of non-photochemical quenching (NPQ), electron transport, and light energy allocation of photosystems in photoprotection needs to be further understood in different light-demanding plants. The activities of photosystem II (PSII) and photosystem I (PSI) in shade-tolerant species Panax notoginseng, intermediate species Polygonatum kingianum, and sun-demanding species Erigeron breviscapus were comparatively measured to elucidate photoprotection mechanisms in different light-demanding plants under dynamic light. The results showed that the NPQ and PSII maximum efficiency (F v\'/F m\') of E. breviscapus were higher than the other two species under dynamic high light. Meanwhile, cyclic electron flow (CEF) of sun plants is larger under transient high light conditions since the slope of post-illumination, P700 dark reduction rate, and plastoquinone (PQ) pool were greater. NPQ was more active and CEF was initiated more readily in shade plants than the two other species under transient light. Moreover, sun plants processed higher quantum yield of PSII photochemistry (ΦPSII), quantum yield of photochemical energy conversion [Y(I)], and quantum yield of non-photochemical energy dissipation due to acceptor side limitation (Y(NA), while the constitutive thermal dissipation and fluorescence (Φf,d) and quantum yield of non-photochemical energy dissipation due to donor side limitation [Y(ND)] of PSI were higher in shade plants. These results suggest that sun plants had higher NPQ and CEF for photoprotection under transient high light and mainly allocated light energy through ΦPSII and ΦNPQ, while shade plants had a higher Φf,d and a larger heat dissipation efficiency of PSI donor. Overall, it has been demonstrated that the photochemical efficiency and photoprotective capacity are greater in sun plants under transient dynamic light, while shade plants are more sensitive to transient dynamic light.

The Yellow Sea green tide (YSGT) is the world\'s largest transregional macroalgal blooms, and the causative species Ulva prolifera (U. prolifera) suffers from ultraviolet-b radiation (UVBR) during the floating migration process. Previous study confirmed that U. prolifera displayed a wide variety of physiological responses characterized as acclimation to UVBR, while the response mechanisms against low-dose and short-term radiation (LDSTR) are not clear. A study with photosynthetically active radiation (PAR) and UVBR was designed: normal light (NL: 72 μmol photons m-2 s-1), NL+0.3 (UVBR: 0.3 W·m-2), and NL+1.6 (UVBR: 1.6 W·m-2). The results showed that high-dose UVBR inhibited photosynthesis in thalli, especially under long-term exposure, while a variety of physiological responses were observed under LDSTR. The inhibition of photosynthesis appeared to be ameliorated by the algae under LDSTR. Further analysis showed that U. prolifera achieved balancing damage by means of non-photochemical quenching (NPQ), accumulation of phenolic compounds coupled with the ASA-GSH cycle involved in the antioxidant process and enhanced photorespiratory metabolism under LDSTR. This study provides new insights into the balancing damage mechanisms of U. prolifera under LDSTR, enabling the thalli to adapt to the light conditions during the long duration and distance involved in floating migration.

Light is essential for photosynthesis but light levels that exceed an organism\'s assimilation capacity can cause serious damage or even cell death. Plants and microalgae have developed photoprotective mechanisms collectively referred to as non-photochemical quenching to minimize such potential damage. One such mechanism is energy-dependent quenching (qE), which dissipates excess light energy as heat. Over the last 30 years, much has been learned about the molecular mechanism of qE in green algae and plants. However, the steps between light perception and qE represented a gap in our knowledge until the recent identification of light-signaling pathways that function in these processes in the green alga Chlamydomonas reinhardtii. In this review, we summarize the high light and UV-mediated signaling pathways for qE in Chlamydomonas. We discuss key questions remaining about the pathway from light perception to photoprotective gene expression in Chlamydomonas. We detail possible differences between green algae and plants in light-signaling mechanisms for qE and emphasize the importance of research on light-signaling mechanisms for qE in plants.

Light is highly heterogeneous in natural conditions, and plants need to evolve a series of strategies to acclimate the dynamic light since it is immobile. The present study aimed to elucidate the response of light reaction of photosynthesis to dynamic sunflecks in a shade-tolerant species Panax notoginseng and to examine the regulatory mechanisms involved in an adaptation to the simulated sunflecks. When P. notoginseng was exposed to the simulated sunflecks, non-photochemical quenching (NPQ) increased rapidly to the maximum value. Moreover, in response to the simulated sunflecks, there was a rapid increase in light-dependent heat dissipation quantum efficiency of photosystem II (PSII) (ΦNPQ), while the maximum quantum yield of PSII under light (F v\'/F m\') declined. The relatively high fluorescence and constitutive heat dissipation quantum efficiency of PSII (Φf,d) in the plants exposed to transient high light (400, 800, and 1,600 μmol m-2 s-1) was accompanied by the low effective photochemical quantum yield of PSII (ΦPSII) after the dark recovery for 15 min, whereas the plants exposed to transient low light (50 μmol m-2 s-1) has been shown to lead to significant elevation in ΦPSII after darkness recovery. Furthermore, PSII fluorescence and constitutive heat dissipation electron transfer rate (J f,d) was increased with the intensity of the simulated sunflecks, the residual absorbed energy used for the non-net carboxylative processes (J NC) was decreased when the response of electron transfer rate of NPQ pathway of PSII (J NPQ) to transient low light is restricted. In addition, the acceptor-side limitation of PSI [Y(NA)] was increased, while the donor-side limitation of photosystems I (PSI) [Y(ND)] was decreased at transient high light conditions accompanied with active cyclic electron flow (CEF). Meanwhile, when the leaves were exposed to transient high light, the xanthophyll cycle (V cycle) was activated and subsequently, the J NPQ began to increase. The de-epoxidation state [(Z + A)/(V + A + Z)] was strongly correlated with NPQ in response to the sunflecks. In the present study, a rapid engagement of lutein epoxide (Lx) after the low intensity of sunfleck together with the lower NPQ contributed to an elevation in the maximum photochemical quantum efficiency of PSII under the light. The analysis based on the correlation between the CEF and electron flow devoted to Ribulose-1, 5-bisphosphate (RuBP) oxygenation (J O) indicated that at a high light intensity of sunflecks, the electron flow largely devoted to RuBP oxygenation would contribute to the operation of the CEF. Overall, photorespiration plays an important role in regulating the CEF of the shade-tolerant species, such as P. notoginseng in response to transient high light, whereas active Lx cycle together with the decelerated NPQ may be an effective mechanism of elevating the maximum photochemical quantum efficiency of PSII under light exposure to transient low light.