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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.)

Differences in plant shade tolerance constitute a major mechanism driving the succession of forest communities in subtropical forests. However, the indirect effects of differences in light requirements on the growth of mid- and late-successional tree species are unclear, and this potential growth effect has not been explained at the transcriptome level. Here, a typical mid-successional dominant tree species, Schima superba Gardn. et Champ, and a typical late-successional dominant tree species, Cryptocarya concinna Hance were used as materials and planted under 100% full light (FL) and 30% FL (low light, LL) to explore the responses of tree species in different successional stages of subtropical forests to different light environments. Transcriptome sequencing was used to analyze the expression changes in genes related to growth and photoprotection under different light environments. The young leaves of S. superba accumulated more malondialdehyde (MDA) and superoxide radicals (${\mathrm{O}}_2^{{{}^{\bullet}}^{-}}$) under LL. A lower hormone content (auxin, cytokinin, gibberellin) in the young leaves, a weaker photosynthetic capacity in the mature leaves and significant downregulation of related gene expression were also found under LL, which resulted in the total biomass of S. superba under LL being lower than that under FL. The young leaves of C. concinna had less MDA and ${\mathrm{O}}_2^{{{}^{\bullet}}^{-}}$, and a higher hormone contents under LL than those under FL. There was no significant difference in photosynthetic capacity between mature leaves in contrasting light environments. Although the biomass of C. concinna under LL was less than that under FL, the height of C. concinna under LL was higher than that under FL, indicating that C. concinna could grow well under the two light environments. Our results describing the acclimatization of light at the physiological, molecular and transcriptome levels are important for a complete understanding of successional mechanisms., (© The Author(s) 2021. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

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:Pgross) 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.

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.)

Light-driven subtropical forest succession is a dynamic process in which mesophytic climax communities replace heliophytic ones. Juvenile leaves (particularly mesophytic ones) are sensitive to high irradiances. To determine the photoprotection strategy that juvenile leaves use during subtropical forest succession, anthocyanin accumulation patterns were investigated in the young leaves of two mid-successional dominant trees (i.e., Schima superba and Castanopsis fissa) and two late-successional dominant trees (i.e., Cryptocarya concinna and Acmena acuminatissima) grown in 100% (FL) and 25% (LL) of full sunlight. All four tree species produced anthocyanins in their juvenile leaves when <50% of chlorophylls and carotenoids had developed. Higher anthocyanin concentrations accumulated in the young leaves grown in FL than in those grown in LL and in late-successional than in mid-successional trees. The juvenile leaves of late-successional trees were subjected to higher light-induced photoinhibition than those of mid-successional trees, despite of the fact that the leaves of late-successional trees showed greater non-photochemical quenching than those of mid-successional trees. Under LL conditions, photosystem II excitation pressure (1 - qP) was significantly higher in the juvenile leaves of late-successional trees than those of mid-successional trees. Under either FL or LL conditions, anthocyanin concentrations in juvenile leaves were negatively related to the light compensation point in mature leaves across species. However, anthocyanin concentrations were positively related to the antioxidant capacity of juvenile leaves. These results indicate that anthocyanin accumulation in the juvenile leaves of subtropical dominant trees during forest community succession is a flexible photoprotective response to ambient irradiances according to leaf sensitivity to light., (© The Author 2016. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

We selected five typical tree species, including one early-successional species (ES) Pinus massoniana Lamb., two mid-successional species (MS) Schima superba Gardn. et Champ. and Castanopsis fissa (Champ. ex Benth.) Rehd. et Wils. and two late-successional species (LS) Cryptocarya concinna Hance. and Acmena acuminatissima (BI.) Merr et Perry., which represent the plants at three successional periods in Dinghushan subtropical forest succession of southern China. Potted seedlings of the five species were grown under 12% of full sunlight for 36 months. The ES and MS showed the slowest and fastest responses to lightflecks, respectively, which correlated with the rate of stomatal opening. In contrast to P. massoniana and C. concinna, the other three species exhibited a high induction loss. Early-successional species showed the lowest specific leaf area and chlorophyll content, the highest photosynthetic capacity (A(max)) and respiratory carbon losses (R(d)). Compared with ES and MS, LS showed lower A(max) and R(d). The five tree species showed a similar chlorophyll a/b ratio after long-term low-light adaptations. On the other hand, LS had a relatively higher de-epoxidation state to protect themselves from excess light during lightflecks. Our results indicated that (i) slower responses to lightflecks could partially explain why ES species could not achieve seedling regeneration in low-light conditions; (ii) fast responses to lightflecks could partially explain why MS species could achieve seedling regeneration in low-light conditions; and (iii) smaller respiratory carbon losses might confer on the LS species a competitive advantage in low-light conditions.