Photosynthetica - Latest articles
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Photosynthetic mechanism of high yield under an improved wide-narrow row planting pattern in maize
X.L. GE, Y.B. CHEN, Y. WANG, B.C. WANG, Q. CHAO, Y. YU, X.J. GONG, Y.B. HAO, L. LI, Y.B. JIANG, G.Y. LV, C.R. QIAN, C.D. JIANG
Photosynthetica 2022, 60(3):465-475 | DOI: 10.32615/ps.2022.032 
Wide-narrow row maize planting patterns are a popular way to enhance maize yield via improving canopy PAR. To further optimize canopy PAR, we designed an improved wide-narrow row planting pattern (R2) based on the principle of the shortest projection length and the longest illumination of objects on the ground. Compared to the traditional wide-narrow row planting pattern (R1), maize yield increased by about 10% in R2. R2 maize had higher PAR, leaf area index, chlorophyll content, and photosynthetic rates than maize grown in R1. Moreover, compared to maize leaves in R1, the carbon assimilation enzymatic activities were also significantly higher...
Diurnal decline in the photosynthetic capacity of uppermost leaves in an eggplant canopy grown in a horticultural greenhouse
K. NOMURA, M. SAITO, M. ITO, S. YAMANE, T. IWAO, I. TADA, T. YAMAZAKI, S. ONO, D. YASUTAKE, M. KITANO
Photosynthetica 2022, 60(3):457-464 | DOI: 10.32615/ps.2022.040
Parameters representing leaf photosynthetic capacity, namely, the maximal carboxylation rate (Vcmax), maximal electron transport rate (Jmax), and triose phosphate-utilization rate (Tp), can vary depending on various factors. The present study investigated diurnal variations in Vcmax, Jmax, and Tp of uppermost leaves of soil-grown, well-watered eggplant in a greenhouse based on the simultaneous measurements of leaf gas exchange and chlorophyll fluorescence. The values of net photosynthetic rates and electron transport rates plotted against intercellular...
Comparative study of photosynthetic capacity in lower leaves in the canopy of dwarf and semidwarf wheat
H.W. LI, J. ZHANG, Q. ZHENG, B. LI, Z.S. LI
Photosynthetica 2022, 60(3):445-456 | DOI: 10.32615/ps.2022.037
Semidwarf wheat contributes to significant yield increase worldwide, however, few dwarf cultivars of wheat are cultivated due to grain yield penalty. In this study, a new dwarf wheat Triticum aestivum L., cv. Xiaoyan 101, was investigated to explore its photosynthetic performance and yield potential. In the comparison of the semidwarf wheat cultivars, Jing 411 and Xiaoyan 101, although the first three leaves (including flag leaves) did not differ significantly in both genotypes, Xiaoyan 101 conferred a higher content of photosynthetic pigments and higher photochemical efficiency but had lower contents of hydrogen peroxide and malondialdehyde...
Ultrastructure, adaptability, and alleviation mechanisms of photosynthetic apparatus in plants under waterlogging: A review
S. SHARMA, U. BHATT, J. SHARMA, H.M. KALAJI, J. MOJSKI, V. SONI
Photosynthetica 2022, 60(3):430-444 | DOI: 10.32615/ps.2022.033
Photosynthesis is a process highly sensitive to various abiotic and biotic stresses in plants. Among them, the major abiotic stress, waterlogging, affects the crop's growth and productivity. Under waterlogging, the photosynthetic apparatus of plants was destroyed. Waterlogging reduced chlorophyll content and the net photosynthetic rate. Therefore, this updated review summarized the effect of waterlogging on chloroplast ultrastructure, photosynthetic characteristics, and chlorophyll fluorescence attributes of plant species. By studying various research papers, we found that intercellular concentration of available carbon dioxide in mesophyll cells,...
The effects of nitrogen application on the growth, photosynthesis, and antioxidant activity of Amaranthus viridis
C.-C. CHEN, M.-Y. HUANG, K.-H. LIN, M.-T. HSUEH
Photosynthetica 2022, 60(3):420-429 | DOI: 10.32615/ps.2022.034
Amaranthus viridis is a functional food due to its antioxidant activity. The aim of this study was to investigate the responses of photosynthesis, growth, and antioxidant properties in A. viridis to nitrogen (N) applications. A. viridis plants were cultivated under low N (LN), medium N (MN), and high N (HN), and harvested at the reproductive phase. The dry mass and plant height of A. viridis plants increased with elevated N, and the dry mass of HN was saturated. Net photosynthetic rate, stomatal conductance, and water-use efficiency in the leaves at HN were strengthened. Meanwhile, under HN, chlorophylls (Chl), their precursors,...
Both uniconazole and 5-aminolevulinic acid increase maize (Zea mays L.) yield by changing its ear morphology and increasing photosynthetic efficiency and antioxidants in saline-alkali land
L. XU, N.J. FENG, X.L. LIANG, H.H. ZHAO, S.Y. WANG, Y. JIANG, Y. ZHAO, D.F. ZHENG
Photosynthetica 2022, 60(3):408-419 | DOI: 10.32615/ps.2022.029
Saline-alkaline stress is one of the most detrimental abiotic stresses that restrict the yield and physiological activity of maize (Zea mays L.). In the present study, maize was planted on saline-alkali land, while 25 mg L-1 uniconazole (S3307) and 40 mg L-1 5-aminolevulinic acid (ALA) were sprayed at the stage of nine expanded leaves. Our results showed that both S3307 and ALA applications significantly increased all ear width, volume, and mass in the maturity stage. Both applications also upregulated photosynthetic efficiency via increasing the chlorophyll content, net photosynthetic rate, transpiration rate, and...
Systematic salt tolerance-related physiological mechanisms of wild soybean and their role in the photosynthetic activity and Na+ distribution of grafted soybean plants
Z.C. XUE, Y. WANG, J. LIU
Photosynthetica 2022, 60(3):400-407 | DOI: 10.32615/ps.2022.030
Systematic salt tolerance-related physiological mechanisms in roots and shoots of halophyte Dongying wild soybean have not yet been thoroughly studied. In this study, photosynthesis, modulated 820-nm reflection, chlorophyll a fluorescence, and Na+ distribution in cultivated (Gmc) and wild (Gsw) soybean leaves of grafted soybean plants were investigated after NaCl treatment. Results showed that the decreases in photosynthetic rate, performance index, active P700 content, and plastocyanin reduction were significantly greater in the Gsw leaves than those in the Gmc...
Mitigative effect of 6-benzyladenine on photosynthetic capacity and leaf ultrastructure of maize seedlings under waterlogging stress
J. WANG, Y.L. WANG, D.Y. WANG, J.X. HUANG, Y.B. LIU, M. ZHU, F.H. LI
Photosynthetica 2022, 60(3):389-399 | DOI: 10.32615/ps.2022.027
6-Benzyladenine (6-BA) is an artificial synthetic cytokinin, which plays an important role in regulating plant responses to abiotic stress. This study aimed to investigate the mitigative effect of exogenous 6-BA on photosynthetic capacities and leaf ultrastructure under waterlogging stress using two waxy corn inbred lines. The results showed that waterlogging stress disrupted the photosynthesis of waxy corn seedlings. However, exogenous 6-BA alleviated the inhibition caused by waterlogging stress. Under the waterlogging conditions, 6-BA treatment of plants helped preserve the structural...
Biochemical, gas exchange, and chlorophyll fluorescence analysis of maize genotypes under drought stress reveals important insights into their interaction and homeostasis
G.M. SINGH, S. GOLDBERG, D. SCHAEFER, F. ZHANG, S. SHARMA, V.K. MISHRA, J. XU
Photosynthetica 2022, 60(3):376-388 | DOI: 10.32615/ps.2022.024
Many studies have been conducted on maize to study the effect of drought on yield at the flowering stage, but understanding biochemical and photosynthetic response against drought at the seedling stage needs to be well established. Thus, to understand differential changes and interaction of biochemical and photosynthetic parameters at the seedling stage under drought, a greenhouse experiment with twelve maize genotypes under severe drought (30% field capacity) and irrigated (90-100% field capacity) conditions were performed. Drought differentially altered biochemical and photosynthetic parameters in all genotypes. A sharp increase in hydrogen peroxide,...
László Mustárdy (1945-2022)
G. GARAB
Photosynthetica 2022, 60(3):372-375 | DOI: 10.32615/ps.2022.028
In photosynthesis research, in the early 1970s, one of the main problems was to understand the structure and functions of the thylakoid membranes of chloroplasts of vascular plants. László Mustárdy, with his great skill and legendary patience, took care excellently of the tasks of electron microscopic ultrastructure.
Comparative PSII photochemistry of quinoa and maize under mild to severe drought stress
C. MALAN, J.M. BERNER
Photosynthetica 2022, 60(3):362-371 | DOI: 10.32615/ps.2022.022
Quinoa has been identified as a climate-resilient crop that can overcome unfavorable conditions. This study explores the photochemical efficiency of quinoa compared to maize subjected to drought stress. The JIP-test was used to assess the photochemical efficiency of both crops. Proline content, leaf water potential, and membrane leakage were also determined. The maximum photochemical efficiency (Fv/Fm) did not change for quinoa and maize under moderate stress. However, severe drought conditions resulted in a decline in Fv/Fm in maize but not quinoa. Furthermore, the PSII performance index (PIABS,total)...
Bacon Ke (1920-2022): a pioneer of primary photochemistry of photosynthesis
G. GOVINDJEE, R. MALKIN, T. OGAWA
Photosynthetica 2022, 60(3):360-361 | DOI: 10.32615/ps.2022.026
Bacon Ke was a physical chemist and an ingenious instrument designer whose work led to an in-depth understanding of the primary photochemistry of photosynthesis.
Speed of light-induced stomatal movement is not correlated to initial or final stomatal conductance in rice
Z. XIONG, Q.W. LUO, D.L. XIONG, K.H. CUI, S.B. PENG, J.L. HUANG
Photosynthetica 2022, 60(3):350-359 | DOI: 10.32615/ps.2022.013
In nature, plants are often confronted with wide variations in light intensity, which may cause a massive carbon loss and water waste. Here, we investigated the response of photosynthetic rate and stomatal conductance to fluctuating light among ten rice genotypes and their influence on plant acclimation and intrinsic water-use efficiency (WUEi). Significant differences were observed in photosynthetic induction and stomatal kinetics across rice genotypes. However, no significant correlation was observed between steady-state and non-steady-state gas exchange. Genotypes with a greater range of steady-state and faster response rate of the gas...
Photosynthetic efficiency of young Rhizophora mangle L. in a mangrove in southeastern Brazil
S.S. PASCOALINI, M.M.P. TOGNELLA, A.R. FALQUETO, M.L.G. SOARES
Photosynthetica 2022, 60(3):337-349 | DOI: 10.32615/ps.2022.025
The study proposes to evaluate the photosynthetic plasticity of Rhizophora mangle L. in four mangrove sites distributed along with the Great Vitória Estuarine System. The variation in organic matter content, which implies the higher essential nutrient availability, contributed to better energy flux performance related to electron transport. Furthermore, salinity damaged the reaction centers (RC), since the site with the highest salinity showed changes in the number and size of active photosynthetic RC and in the specific energy flows per active RC (absorption flux, trapped energy flux, and dissipated energy flux), but the plasticity of the species...
Diurnal photosynthetic performance of two oak species from two provenances in a Mediterranean and a central European common garden
V. HOLLAND, N. REININGER, F. BANTIS, W. BRÜGGEMANN, K. RADOGLOU, M.N. FOTELLI
Photosynthetica 2022, 60(3):326-336 | DOI: 10.32615/ps.2022.023
Oaks may contribute to the stabilization of European forests under climate change. We utilized two common gardens established in contrasting growth regimes, in Greece (Olympiada) and Germany (Schwanheim), to compare the diurnal photosynthetic performance of a Greek and an Italian provenance of two Mediterranean oaks (Quercus pubescens and Q. frainetto) during the 2019 growing season. Although the higher radiation in the southern common garden led to a strong midday depression of chlorophyll a fluorescence parameters (maximum quantum efficiency of PSII, performance index on absorption basis), comparable light-saturated net photosynthetic...
Tribute to Jean Lavorel (1928-2021), an outstanding experimenter and a brilliant theorist of photosynthesis
G. GOVINDJEE, Y. DE KOUCHKOVSKY, G. GARAB
Photosynthetica 2022, 60(2):323-325 | DOI: 10.32615/ps.2022.021
Tribute to Jean Lavorel (16 March 1928–12 January 2021), a pioneer of the ‘Light Reactions of Photosynthesis’. He was known not only for his ingenuity in devising new instruments but in thoroughly analyzing all the available data theoretically and mathematically – mostly all by himself. He measured, elegantly, oxygen evolution and light given off by photosynthetic organisms, both prompt and delayed chlorophyll fluorescence. He ingeniously used these data to understand how light energy is converted to chemical energy in natural systems. We present below a summary of his life and research.
Updating the occurrence of crassulacean acid metabolism (CAM) in the genus Clusia through carbon isotope analysis of species from Colombia
P. PACHON, K. WINTER, E. LASSO
Photosynthetica 2022, 60(2):304-322 | DOI: 10.32615/ps.2022.018
Clusia is a widely distributed neotropical genus with 321 currently described species. This remarkable genus is the only one known to contain trees sensu stricto with CAM photosynthesis. To survey the occurrence of CAM in Clusia species from Colombia, we determined the leaf stable carbon isotope composition (δ13C) of 568 specimens from 114 species deposited in 12 Colombian herbaria. In the vast majority of specimens, δ13C values indicated that C3 photosynthesis was the principal contributor to carbon gain. δ13C values typical of strong CAM (less negative than -20‰)...
Effect of application of iron (Fe) and α-ketoglutaric acid on growth, photosynthesis, and Fe content in fragrant rice seedlings
S.R. CHENG, S.S. LI, Z.W. LIANG, F.C. HUANG, X.Q. WU, Z.Y. HAN, X.B. HUANG, X.M. HUANG, Y. REN
Photosynthetica 2022, 60(2):293-303 | DOI: 10.32615/ps.2022.020
At a three-leaf stage, two Fe treatments [0 mg kg-1 (Fe-) and 20 mg.kg-1 (Fe+) in the form of FeCl3] were used in the soil of the pot and then two concentrations of α-ketoglutaric acid [0 mg L-1 (A-) and 50 mg L-1 (A+)] were sprayed to the rice plants of Meixiangzhan and Yuxiangyouzhan cultivars. We showed that seedlings exhibited an increased length and fresh and dry mass of shoots and roots with treatments Fe+A- and Fe-A+, as well as the Fe content increased greatly. Both treatments increased the morphological characteristic values of roots and promoted photosynthesis. Interestingly, Fe+A+...
Response of upland cotton (Gossypium hirsutum L.) leaf chlorophyll content to high heat and low-soil water in the Arizona low desert
A.L. THOMPSON, M.M. CONLEY, M.T. HERRITT, K.R. THORP
Photosynthetica 2022, 60(2):280-292 | DOI: 10.32615/ps.2022.014
The sustainable production of upland cotton, an economically important fiber crop, is threatened by changing environmental factors including high temperatures and low-soil water content. Both high heat and low-soil water can reduce net photosynthesis resulting in low fiber yields or poor fiber quality. Leaf chlorophyll content has a direct relationship with photosynthetic rate. Understanding how high heat and low-soil water affect chlorophyll content can identify opportunities for breeding improvement that will lead to sustainable fiber yields. A two-year field trial located in Maricopa Arizona measured leaf chlorophyll content, available soil water,...
Upregulation of the mitochondrial alternative oxidase pathway improves PSII function and photosynthetic electron transport in tomato seedlings under chilling stress
J.J. ZENG, W.H. HU, X.H. HU, H.M. TAO, L. ZHONG, L.L. LIU
Photosynthetica 2022, 60(2):271-279 | DOI: 10.32615/ps.2022.019
The aim of this study was to explore how the mitochondrial alternative oxidase (AOX) pathway alleviates photoinhibition in chilled tomato (Solanum lycopersicum) seedlings. Chilling induced photoinhibition in tomato seedlings despite the increases in thermal energy dissipation and cyclic electron flow around PSI (CEF-PSI). Chilling inhibited the function of PSII and blocked electron transport at the PSII acceptor side, however, it did not affect the oxygen-evolving complex on the donor side of PSII. Upregulation of the AOX pathway protects against photoinhibition by improving PSII function and photosynthetic electron transport in tomato seedlings...
The role of photosynthetic activity in the regulation of flg22-induced local and systemic defence reaction in tomato
Z. CZÉKUS, P. KOPRIVANACZ, A. KUKRI, N. IQBAL, A. ÖRDÖG, P. POÓR
Photosynthetica 2022, 60(2):259-270 | DOI: 10.32615/ps.2022.015
Flagellin (flg22) induces rapid and long-lasting defence responses. It may also affect the photosynthetic activity depending on several internal and external factors, such as the phytohormone ethylene or the day/night time. Based on the results, flg22 treatment, neither in the light phase nor in the evening, caused any significant change in chlorophyll fluorescence induction parameters in the leaves of wild-type and ethylene-receptor mutant Never ripe tomato plants measured the next morning. However, flg22 in the light phase decreased the effective quantum yield and the photochemical quenching both locally and systemically in guard cells. In...
Vertical profile of photosynthetic CO2 response within rice canopy
Y.P. LV, Y. PAN
Photosynthetica 2022, 60(2):251-258 | DOI: 10.32615/ps.2022.011
Leaf-specific Farquhar-von Caemmerer-Berry (FvCB) model was fitted to characterize the vertical profile of photosynthetic CO2 response within rice canopy. Leaf-position-specific and canopy average FvCB models were fitted to study a suitable leaf representing photosynthetic parameters at the canopy scale. The results showed that leaf photosynthesis was limited by Rubisco activity or ribulose-1,5-bisphosphate regeneration under field conditions. The maximum rate of carboxylation, maximum rate of electron transport, rate of triose phosphates utilization, and light respiration rate in the FvCB model reached the highest values for the top second...
Leaf structure and photosynthesis in Populus alba under naturally fluctuating environments
X.Y. LIN, X.X. WANG, Q.Y. ZENG, Q. YANG
Photosynthetica 2022, 60(2):240-250 | DOI: 10.32615/ps.2022.012
The ability to modulate photosynthesis is essential for plants to adapt to fluctuating growing conditions. Populus species show high tolerance to various and highly variable environments. To understand their response strategies against fluctuating environments, this study investigated the morphological and physiological differences of white poplar (Populus alba) leaves when grown in a phytotron, glasshouse, and field. Our results show that the palisade cells were elongated in the field, which would enhance intercellular CO2 exchange. Photosynthetic capacity was the highest in the field leaves, as shown by higher electron transport...
Chlorophyll measurements in Alstroemeria sp. using SPAD-502 meterand the color space CIE L*a*b*, and its validation in foliar senescence
I. VILLEGAS-VELÁZQUEZ, H.A. ZAVALETA-MANCERA, M.L. ARÉVALO-GALARZA, J. SUAREZ-ESPINOSA, C. GARCIA-OSORIO, D. PADILLA-CHACON, I.G. GALVAN-ESCOBEDO, J.F. JIMENEZ-BREMONT
Photosynthetica 2022, 60(2):230-239 | DOI: 10.32615/ps.2022.007
Our research aimed to study the correlation between the SPAD-502 readings and the color space CIE L*a*b* values in two cultivars of Alstroemeria sp. during leaf senescence and to evaluate the statistical criteria used in the selection of the best fit calibration functions. We demonstrate the importance of the Akaike information criterion and the parsimonious function besides the coefficient of determination. The reliability of the functions was tested by Student's t-test comparison between the chlorophyll (Chl) estimated from SPAD readings and their chemical concentrations. Polynomial and Hoerl function described well the...
Transgenerational effect of UV-B priming on photochemistry and associated metabolism in rice seedlings subjected to PEG-induced osmotic stress
A. SEN, J.T. PUTHUR, D. CHALLABATHULA, M. BRESTIČ
Photosynthetica 2022, 60(2):219-229 | DOI: 10.32615/ps.2022.006
Rice being the major food crop for more than half of the world population is severely affected by drought stress starting from the establishment of the seedling. We focused on the UV-B priming mediated transgenerational drought tolerance of a drought-tolerant rice variety (Vaisakh) towards polyethylene glycol (PEG) 6,000 (20%)-induced drought. Results showed that priming in F0 generation and re-priming in F1 generation with UV-B enhanced the PEG stress tolerance potential of rice seedlings with increased expression of genes encoding antioxidant enzymes and stress-related proteins offering better protection to primed plants. UV-B...
Chloroplast protease/chaperone AtDeg2 holds γ1 subunit of ATP synthase in an unaggregated state under high irradiance conditions in Arabidopsis thaliana
P. JAGODZIK, G. JACKOWSKI
Photosynthetica 2022, 60(2):212-218 | DOI: 10.32615/ps.2022.004
Little data on the role played in vivo by chloroplast protein AtDeg2 as a chaperone is available. Therefore, we sought for chloroplast proteins protected from high irradiance-induced interprotein aggregation via disulphide bridges by AtDeg2 acting as a holdase. To reach this goal, we performed analyses which involved comparative diagonal electrophoreses of lysates of chloroplasts isolated from wild type (WT) plants and transgenic plants 35S:AtDEG2ΔPDZ1-GFP which expressed AtDeg2 lacking its chaperone activity but retaining the protease activity. The results of the analyses indicate that AtDeg2 acting...
Effect of short-term combined alkaline stress on antioxidant metabolism, photosynthesis, and leaf-air temperature difference in sorghum
J.J. GUO, X.X. XU, R.D. ZHANG, X.F. CHEN, Y.F. XING, B. LI, C. LIU, Y.F. ZHOU
Photosynthetica 2022, 60(2):200-211 | DOI: 10.32615/ps.2021.067
Alkaline stress is important abiotic stress that restricts the growth and physiological activity of sorghum (Sorghum bicolor L. Moench). We aimed to investigate the effects of alkaline stress on alkali-tolerant SX44B and alkali-sensitive 262B sorghum inbred lines. The results showed that alkaline stress decreased the content of chlorophyll, activity of photosystem II, net photosynthetic rate, and destroyed chloroplast morphology. These changes were less pronounced in SX44B, possibly owing to its higher antioxidant enzyme activity and nonphotochemical quenching. Alkaline stress decreased water content, transpiration rate, and stomatal conductance...
Effects of source-sink alteration by pruning on physiological parameters and fruit production of Rosa roxburghii Tratt. on the Yunnan-Guizhou Plateau in China
C. ZHANG, K. YAN, L.Z. LIN, Y.M. FANG, X.Y. ZHANG
Photosynthetica 2022, 60(2):190-199 | DOI: 10.32615/ps.2022.001
The formation of economic yield and fruit quality of Roxburgh rose (Rosa roxburghii Tratt.) depends essentially on its source-sink interaction. Thus, a pruning experiment was conducted to assess the effects of source-sink regulation on photosynthetic physiology and fruit yield of Roxburgh rose, which was of great significance to production. Cutting off vegetative branches reduced physiological fruit loss and malformed fruits but increased single fruit quality and yield. Results revealed that the stomatal characteristics, the composition of mesophyll tissue, and photosynthesis of leaves on reproductive branches were significantly affected by...
Nitrogen rate and plant density interaction enhances grain yield by regulating the grain distribution of secondary branches on the panicle axis and photosynthesis in japonica rice
Y.L. GONG, Y. LEI, X.P. ZHANG, B.C. YAN, X.T. JU, X.Y. CHENG, J.D. ZHANG, X.Y. SUN, H. XU, W.F. CHEN
Photosynthetica 2022, 60(2):179-189 | DOI: 10.32615/ps.2022.002
Two japonica rice cultivars with different panicle trait index (PTI), HP917 (a high-PTI cultivar) and DP128 (a low-PTI cultivar) were used to investigate the effects of the nitrogen (N) rate and plant density on the grain distribution of secondary branches on the panicle axis, leaf photosynthetic characteristics, and grain yield by a split plot design. The main plots were assigned to four N rates (0, 140, 200, and 260 kg ha-1), and the subplots were assigned to two plant densities: (D20, 15 plants m-2; D10, 30 plants m-2). Results showed that the grain yield was increased by increasing N rate...
Partial shade improved the photosynthetic capacity and polysaccharide accumulation of the medicinal plant Bletilla ochracea Schltr.
X.F. YU, X.Y. MING, M. XIONG, C. ZHANG, L.J. YUE, L. YANG, C.Y. FAN
Photosynthetica 2022, 60(2):168-178 | DOI: 10.32615/ps.2021.064
To study the light intensity suitable for Bletilla ochracea Schltr., morphology, photosynthetic parameters, and polysaccharide content of seedlings were evaluated under different light intensities. All shade treatments promoted plant growth and net photosynthetic rate while having no significant effect on transpiration rate. The maximum photochemical efficiency and potential photochemical efficiency reached the lowest values under full sunlight. The electron transport rate and photochemical quenching under shade were significantly higher than those under full light, while nonphotochemical quenching was the highest under full light. This indicated...