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Many studies have investigated the photoprotective and photosynthetic capacity of plant leaves, but few have simultaneously evaluated the dynamic changes of photoprotective capacity and photosynthetic maturation of leaves at different developmental stages. As a result, the process between the decline of photoprotective substances and the onset of photosynthetic maturation during plant leaf development are still poorly understood, and the relationship between them has not been quantitatively described. In this study, the contents of photoprotective substances, photosynthetic pigment content and photosynthetic capacity of leaves at different developmental stages from young leaves to mature leaves were determined by spatio-temporal replacement in eight dominant tree species in subtropical evergreen broadleaved forests. The correlation analysis found that the data sets of anthocyanins, flavonoids, total phenolics and total antioxidant capacity were mainly distributed on one side of the symmetry axis (y = x), while the data sets of flavonoids, total phenolics and total antioxidant capacity were mainly distributed on both sides of the symmetry axis (y = x). In addition, the content of photoprotective substances in plant leaves was significantly negatively correlated with photosynthetic pigment content and photosynthetic capacity but was significantly positively correlated with dark respiration rate (Rd). When chlorophyll accumulated to ~50% of the final value, the photoprotective substance content and Rd of plant leaves reached the lowest level, and anthocyanins disappeared completely; in contrast, the photosynthetic capacity reached the highest level. Our results suggest that anthocyanins mainly play a light-shielding role in the young leaves of most plants in subtropical forests. In addition, 50% chlorophyll accumulation in most plant leaves was the basis for judging leaf photosynthetic maturity. We also believe that 50% chlorophyll accumulation is a critical period in the transition of plant leaves from high photoprotective capacity (high metabolic capacity, low photosynthetic capacity) to low photoprotective capacity (low metabolic capacity, high photosynthetic capacity)., (© The Author(s) 2023. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Increasing amounts of experimental evidence show that anthocyanins provide physiological protection to plants under stress. However, the difference in photoprotection mediated by anthocyanins and other photoprotective substances in different seasons is still uncertain. To determine the relationship between anthocyanin accumulation and the photoprotective effects in different seasons, Castanopsis chinensis and Acmena acuminatissima, whose anthocyanin accumulation patterns differ in different seasons, were used as materials to explain how plants adapt to different seasons; as such, their physiological and biochemical responses were analyzed. Young leaves of C. chinensis and A. acuminatissima presented different colors in the different seasons. In summer, the young leaves of C. chinensis were purplish red, while those of A. acuminatissima were light green. In winter, the young leaves of C. chinensis were light green, while those of A. acuminatissima were red. Compared with the young red leaves, the young light green leaves that did not accumulate anthocyanins had higher flavonoid and phenolics contents, total antioxidant capacity, non-photochemical quenching (NPQ), and relative membrane leakage, and a slower recovery rate in the maximum photochemical efficiency (F v /F m ) after high-light treatment. In addition, the net photosynthesis rate (P n ), transpiration rate (T r ), stomatal conductance (g s ), and the effective quantum yield of PSII (Φ PSII ) of the young leaves in winter were significantly lower than those in summer, while the activities of catalase (CAT, EC 1.11.1.6), peroxidase (POD, EC 1.11.1.7), and superoxide dismutase (SOD, EC 1.15.1.1) were significantly higher than those in summer. These data indicate that to adapt to seasonal changes anthocyanins, other antioxidative substances and antioxidative enzymes, as well as components involved in the safe dissipation of excitation energy as heat need to cooperate with one another., (© 2020. Springer Nature B.V.)

Anthocyanins are water-soluble pigments in plants known for their photoprotective role against photoinhibitory and photooxidative damage under high light (HL). However, it remains unclear whether light-shielding or antioxidant activity plays a major role in the photoprotection exerted by anthocyanins under HL stress. To shed light on this question, we analyzed the physiological and biochemical responses to HL of three Arabidopsis thaliana lines (Col, chi, ans) with different light absorption and antioxidant characteristics. Under HL, ans had the highest antioxidant capacity, followed by Col, and finally chi; Col had the strongest light attenuation capacity, followed by chi, and finally ans. The line ans had weaker physiological activity of chloroplasts and more severe oxidative damage than chi after HL treatment. Col with highest photoprotection of light absorption capacity had highest resistance to HL among the three lines. The line ans with high antioxidant capacity could not compensate for its disadvantages in HL caused by the absence of the light-shielding function of anthocyanins. In addition, the expression level of the Anthocyanin Synthase (ANS) gene was most upregulated after HL treatment, suggesting that the conversion of colorless into colored anthocyanin precursors was necessary under HL. The contribution of anthocyanins to flavonoids, phenols, and antioxidant capacity increased in the late period of HL, suggesting that plants prefer to synthesize red anthocyanins (a group of colored antioxidants) over other colorless antioxidants to cope with HL. These experimental observations indicate that the light attenuation role of anthocyanins is more important than their antioxidant role in photoprotection., (© 2020. Springer Nature B.V.)

The presence of anthocyanins in young leaves plays an important role in mitigation against photodamage and allows leaves to grow and develop normally. Many studies have reported that foliar anthocyanins are distributed within the vacuoles of mesophyll cells, so we explored the novel defence style of anthocyanin-coated young leaves of Castanopsis fissa, a dominant subtropical forest tree species, via removable trichomes. Anthocyanins were distributed in C. fissa leaf trichomes, which produced a red coating for the young leaves. As young leaves developed and then matured, the thickness and density of the anthocyanin trichomes progressively decreased, the coating finally disappearing, allowing greater utilization of light by mature leaves. In addition to anthocyanins, the trichomes contained a remarkably high amount of phenolics, which enable the red coating to be more efficient in screening ultraviolet light. Compared with mature leaves, the young leaves exhibited lower photosynthetic ability, which was attributable to the reduced chlorophyll and Rubisco contents. Removal of the red coating had little effect on the photosynthetic capacity of young leaves. However, the young leaves without the coating suffered greater light-induced photoinhibition due to greater excess light entering the chloroplast and the production of H 2 O 2 Our results suggest that the anthocyanin coating is photoprotective and this anthocyanin defence style may be a metabolically cost-effective way of adjusting the anthocyanin content in response to demand., (© The Author 2016. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Ten anthocyanin components have been detected in roots of purple sweet potato (Ipomoea batatas Lam.) by high-performance liquid chromatography coupled to diode array detection and electrospray ionization tandem mass spectrometry. All the anthocyanins were exclusively cyanidins or peonidin 3-sophoroside-5-glucosides and their acylated derivatives. The total anthocyanin content in purple sweet potato powder obtained by solid-phase extraction was 66 mg g(-1). A strong capacity of purple sweet potato anthocyanins (PSPA) to scavenge reactive oxygen species (superoxide, hydroxyl radical) and the stable 1,1-diphenyl-2-picrylhydrazyl organic free radical was found in vitro using the electron spin resonance technique. To determine the functional roles of anthocyanins in leaves in vivo, for the first time, supplemental anthocyanins were infiltrated into leaves of Arabidopsis thaliana double mutant of the ecotype Landsberg erecta (tt3tt4) deficient in anthocyanin biosynthesis. Chlorophyll fluorescence imaging showed that anthocyanins significantly ameliorated the inactivation of photosystems II during prolonged high-light (1300 micromol m(-2) s(-1)) exposure. Comet assay of DNA revealed an obvious role of supplemental PSPA in alleviating DNA damage by high light in leaves. Our results suggest that anthocyanins could function in vitro and in vivo to alleviate the direct or indirect oxidative damage of the photosynthetic apparatus and DNA in plants caused by high-light stress.

Light attenuation and antioxidation are the main mechanisms of photoprotection by anthocyanin under high light (HL) stress. Anthocyanin synthase (ANS) is the key enzyme in the downstream portion of anthocyanin synthetic pathways. To explore the role of ANS in photoprotection by anthocyanin under HL stress, homozygous ANS -deficient Arabidopsis mutants were screened from SALK_073183 and SALK_028793. Here, we obtained two deficient mutants, ans - 1 and ans - 2 , which had ANS gene expression levels equal to 5.9 and 32.9% of that of Col respectively. By analysing their physiological and biochemical responses to HL stress, we found that there were positive correlations among ANS expression level, anthocyanin content and resistance to HL. The line with the lowest ANS expression level, ans - 1 , was also the most sensitive to HL, showing the lowest anthocyanin content, chlorophyll content, F v/ F m ratio, and Rubisco content and the highest O2•− accumulation and membrane leakage rate, although it also had the highest antioxidant capacity. Experimental evidence suggests that ANS mainly regulated the light-attenuating function of anthocyanin in photoprotection under HL. Blocking excess light is an important function of anthocyanin that protects plants from HL stress, and a high antioxidant capacity cannot compensate for the absence of the light-shielding function of anthocyanin. Upon high light stress, light-shielding and antioxidation are the main mechanisms of photoprotection by anthocyanin, the final key enzyme in whose synthetic pathways is anthocyanin synthase (ANS). This investigation tackled the role of ANS in photoprotection: ANS mainly regulates the light-shielding of anthocyanin in photoprotection. Additionally, we found that light-shielding is the important function in photoprotection of anthocyanins, and high antioxidant capacity cannot compensate for the absence of the light-shielding. [ABSTRACT FROM AUTHOR]

The abundance of phenolic compounds (including anthocyanins) in leaves is associated with photosynthetic performance, but the regulatory mechanism is unclear. Schima superba Gardn. et Champ. and Cryptocarya concinna Hance. which exhibit distinct anthocyanin accumulation patterns, are dominant tree species in the early- and late-successional stages, respectively, of subtropical forests in China. RNA-seq and analyses of phenolic concentrations, antioxidant capacity and photosynthetic characteristics were performed on young and mature leaves of these two species under contrasting light conditions. The high-light-acclimated young leaves of S. superba and C. concinna and low-light-acclimated young leaves of C. concinna were red. These red leaves had higher ratios of electron transport rate to gross photosynthesis (ETR: P gross) and total antioxidant capacity to chlorophyll (TAC:Chl) than did the green leaves, regardless of light conditions. In addition, the red leaves had a higher expression level of the UDP-glucose:flavonoid 3-O-glucosyltransferase (UFGT) gene than did the green leaves, irrespective of light conditions. Total antioxidant capacity was positively correlated with flavonoid content in C. concinna leaves and with total phenolic content in leaves of both species under both high and low light. Consistent with the measurements of photosynthetic performance and flavonoids:Chl ratio, photosynthesis-related genes were extensively downregulated and flavonoid-pathway-related genes were extensively upregulated in young leaves relative to mature leaves. Under high and low light, both non-photochemical quenching and TAC:Chl, which serve as different types of photoprotective tools, were enhanced in young leaves of S. superba, whereas only TAC:Chl was enhanced in young leaves of C. concinna. Our results indicate that the biosynthesis of phenolic compounds in young leaves is likely enhanced by an imbalance between photosynthetic electron supply and demand and that flavonoids play a larger role in meditating photoprotection in late-successional species than in early-successional ones. [ABSTRACT FROM AUTHOR]

Chlorophyll fluorescence and antioxidative capability in detached leaves of the wild type Arabidopsis thaliana L. ecotype Landsberg erecta ( Ler) and three mutants deficient in anthocyanins biosynthesis ( tt3, tt4, and tt3tt4) were investigated during treatment with temperatures ranging 25–45 °C. In comparison with the wild type, chlorophyll fluorescence parameters Fv/Fm, ΦPSII, electron transport rate (ETR), Fv/Fo and qP in three anthocyanin-deficient mutants showed a more rapidly decreasing rate when the temperature was over 35 °C. Non-photochemical quenching (NPQ) in these mutants was almost completely lost at 44 °C, whereas the content of heat stable protein dropped and the rate of the membrane leakage increased. Fo-temperature curves were obtained by monitoring Fo levels with gradually elevated temperatures from 22 °C to 72 °C at 0.5 °C/min. The inflexion temperatures of Fo were 45.8 °C in Ler, 45.1 °C in tt3, 44.1 °C in tt4 and 42.3 °C in tt3tt4, respectively. The temperatures of maximal Fo in three mutants were 1.9–3.8 °C lower than the wild type plants. Meanwhile, three mutants had lower activities of superoxide dismutase (SOD) and ascorbate peroxidase (APX) and an inferior scavenging capability to DPPH (1.1-diphenyl-2-picrylhy.drazyl) radical under heat stress, and in particular tt3tt4 had the lowest antioxidative potential. The results of the diaminobenzidine-H2O2 histochemical staining showed that H2O2 was accumulated in the leaf vein and mesophyll cells of mutants under treatment at 40 °C, and it was significantly presented in leaf cells of tt3tt4. The sensitivity of Arabidopsis anthocyanins-deficient mutants to high temperatures has revealed that anthocyanins in normal plants might provide protection from high temperature injury, by enhancing its antioxidative capability under high temperature stress. [ABSTRACT FROM AUTHOR]

The leaves of many plants are red during particular stages of their lives, but the adaptive significance of leaf colouration is not yet clearly understood. In order to reveal whether anthocyanins play a similar role (i.e. antioxidants) in different seasonal contexts, this study investigated species with red young leaves in the subtropical forest of Dinghushan biosphere reserve (South China) during summer and winter and compared group leaf characteristics between the two seasons. Of 62 total species, 33 exhibited red young leaves in summer only, 6 in winter only, and 23 in both seasons. The anthocyanins extracted from most of these species had an absorption peak at ~530 nm. Frequency distribution analysis showed that the species containing anthocyanins at levels ranging from 0.02 to 0.04 μmol cm−2 occurred most frequently in summer or winter. Based on conditional grouping of the species, no significant variation was observed in the average anthocyanin contents and antioxidant abilities between summer and winter; the flavonoid content in summer was 2-fold that in winter, whereas the anthocyanin:flavonoid ratio in summer was only half that in winter. Moreover, a positive correlation between anthocyanins and flavonoids was found in summer. Therefore, it is less likely for anthocyanins to serve as antioxidants in summer than winter, because such a function in summer leaves is readily replaced by other flavonoids. [ABSTRACT FROM AUTHOR]

Photoprotection strategies that have evolved in plants to cope with high light (HL) stress provide plants with the ability to resist HL. However, it has not been clearly confirmed which photoprotection strategy is the major HL resistance mechanism. To reveal the major photoprotection mechanism against short-term high light (STHL), the physiological and biochemical responses of three Arabidopsis mutants (Col , chi and ans) under STHL were analyzed in this study. After STHL treatment, the most serious photosynthetic pigment damage was observed in chi plants. At the same time, the degrees of membrane and Rubisco damage in chi was the highest, followed by Col , and ans was the smallest. The results showed that ans with high antioxidant capacity showed higher resistance to STHL treatment than Col containing anthocyanins, while chi with no anthocyanin accumulation and small antioxidant capacity had the lowest resistance. In addition, the gene expression results showed that plants tend to synthesize anthocyanin precursor flavonoids with antioxidant capacity under STHL stress. To further determine the major mechanism of photoprotection under STHL, we also analyzed Arabidopsis lines (Col , CHS1 , CHS2 and tt4) that had the same anthocyanin content but different antioxidant capacities. It was found that CHS2 with high antioxidant capacity had higher cell viability, smaller maximal quantum yield of PSII photochemistry (F v /F m) reduction and less reactive oxygen species (ROS) accumulation under HL treatment of their mesophyll protoplasts. Therefore, the antioxidant capacity provided by antioxidant substances was the major mechanism of plant photoprotection under STHL treatment. • The antioxidant capacity was more important than light shielding effect under STHL. • Plants tend to synthesize anthocyanin precursor flavonoids with antioxidant capacity under STHL. • A higher antioxidant capacity correlated with a higher resistance index. • A higher antioxidant capacity correlated with a higher the mesophyll protoplast survival. [ABSTRACT FROM AUTHOR]

Previous studies have suggested that high light (HL) stress causes photoinhibition in plants, while anthocyanins could protect the photosynthetic apparatus against photoinhibition. However, the photoprotection mechanism of anthocyanins is still ambiguous. We studied physiological responses and molecular changes for CHS -overexpression lines ( CHS1 , CHS2 , CHS3 ), Arabidopsis thaliana ecotype Columbia ( Col ), and T-DNA insertion lines of CHS ( tt4 ) under HL (200 μmol m −2  s −1 ) to explore the photoprotection mechanism of anthocyanins. The results showed that HL induced anthocyanin synthesis and accumulation. The leaves of CHS -overexpression lines turned reddest and the genes, including CHS, DFR, ANS, were expressed at highest levels. Thus, the CHS -overexpression lines maintained the highest photosynthetic capacity and suffered the least damage from HL of the three phenotypes. However, the CHS enzyme and anthocyanins were undetectable in tt4 during the experiment. Correspondingly, chlorophyll fluorescence parameters of tt4 declined greatly. The photosynthetic apparatus and cell membranes were also impaired dramatically. The physiological characteristics of Col were compared between CHS -overexpression lines and tt4 . Together, the results suggest that over-expression of CHS gene enhances HL resistance by synthesizing more anthocyanins, that anthocyanins enhance the adaptability of plants to HL and that they maintain photosynthetic capacity via both antioxidation and attenuation of light. [ABSTRACT FROM AUTHOR]

• Different photoprotection strategies for mid- and late-successional tree species. • Mid- and late-successional tree species response patterns to high-light were different. • The mid-successional tree species better tolerate high-light. • Late-successional tree species has greater restoration ability after high-light stress. Light adaptability is a major driver of succession and evolution in natural forest plant communities. However, the relationship between plants and light is contradictory; light provides energy for plant growth, but excessive light negatively affects plant growth. To reveal the pattern of plant adaptation to diurnal variation in light, photoprotection strategies were analyzed in young and mature leaves of six dominant species at two successional stages. Mature leaves of both mid- and late-successional dominant species exhibited a higher net photosynthetic rate (Pn) than young leaves. In particular, the mature leaves of the mid-successional tree species had higher high-light tolerance because of a higher Pn, water use efficiency (WUE) and electron transfer rate (ETR). In contrast, the mature leaves of the late-successional tree species had a higher restoration ability after high-light stress because of higher thermal energy dissipation and faster stomatal regulation. Young leaves of both the mid- and late-successional dominant species exhibited a higher photoprotection capability than mature leaves. The young leaves of the mid-successional tree species had higher high-light tolerance because of their highest WUE and anthocyanin/chlorophyll (Anth/Chl) and higher flavonoid/chlorophyll (Flav/Chl), total phenolics/chlorophyll (Phen/Chl) and total antioxidant capacity/chlorophyll (TAC/Chl) ratios. The young leaves of the late-successional tree species had a stronger restoration ability due to highest Flav/Chl, Phen/Chl, and TAC/Chl ratios and a higher moisture content and non-photochemical quenching coefficient (qN, NPQ). This study shows that in a high-light environment, when leaves are at the same developmental stage, mid-successional tree species better tolerate high-light, while late-successional tree species have a greater restoration ability after high-light stress. [ABSTRACT FROM AUTHOR]