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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...

Phosphate fertilizers increase CO2 assimilation and yield of soybean in a shaded environment

W. ZHAO, B. ZHENG, T.H. REN, X.H. ZHANG, T.Y. NING, G. LI

Photosynthetica 2022, 60(2):157-167 | DOI: 10.32615/ps.2021.063  

Two light treatments [ambient sunlight (L1) during the entire growth period and 40% shade (L2) from 40 d after sowing until 24 d after flowering] and two phosphate fertilizer treatments [no phosphate fertilizer application (P0) and a conventional phosphate fertilizer application (P1)] were used to determine how phosphate fertilizer regulates soybean [Glycine max (L.) Merr.] photosynthesis under shading. We showed that phosphorus significantly increased chlorophyll content and grain yield under shading. The light-saturated net photosynthetic rate, apparent quantum yield, maximum electron transport rate, and maximum Rubisco carboxylation rate...

Dependence of the rate-limiting steps in the dark-to-light transition of photosystem II on the lipidic environment of the reaction center

M. MAGYAR, P. AKHTAR, G. SIPKA, W. HAN, X. LI, G. HAN, J.-R. SHEN, P.H. LAMBREV, G. GARAB

Photosynthetica 2022, 60(1):147-156 | DOI: 10.32615/ps.2022.016  

In our earlier works, we have identified rate-limiting steps in the dark-to-light transition of PSII. By measuring chlorophyll a fluorescence transients elicited by single-turnover saturating flashes (STSFs) we have shown that in diuron-treated samples an STSF generates only F1 (< Fm) fluorescence level, and to produce the maximum (Fm) level, additional excitations are required, which, however, can only be effective if sufficiently long Δτ waiting times are allowed between the excitations. Biological variations in the half-rise time (Δτ1/2) of the fluorescence increment...

Limiting steps and the contribution of alternative electron flow pathways in the recovery of the photosynthetic functions after freezing-induced desiccation of Haberlea rhodopensis

K. GEORGIEVA, A.V. POPOVA, G. MIHAILOVA, A.G. IVANOV, M. VELITCHKOVA

Photosynthetica 2022, 60(1):136-146 | DOI: 10.32615/ps.2022.008  

Haberlea rhodopensis Friv. is unique with its ability to survive desiccation to an air-dry state during periods of extreme drought and freezing temperatures. To understand its survival strategies, it is important to examine the protective mechanisms not only during desiccation but also during rehydration. We investigated the involvement of alternative cyclic electron pathways during the recovery of photosynthetic functions after freezing-induced desiccation. Using electron transport inhibitors, the role of PGR5-dependent and NDH-dependent PSI-cyclic electron flows and plastid terminal oxidase were assessed during rehydration of desiccated leaves. Recovery...

Photosystem II in bio-photovoltaic devices

R.A. VOLOSHIN, S.M. SHUMILOVA, E.V. ZADNEPROVSKAYA, S.K. ZHARMUKHAMEDOV, S. ALWASEL, H.J.M. HOU, S.I. ALLAKHVERDIEV

Photosynthetica 2022, 60(1):121-135 | DOI: 10.32615/ps.2022.010  

Hybrid photoelectrodes containing biological pigment-protein complexes can be used for environmentally friendly solar energy conversion, herbicide detection, and other applications. The total number of scientific publications on hybrid bio-based devices has grown rapidly over the past decades. Particular attention is paid to the integration of the complexes of PSII into photoelectrochemical devices. A notable feature of these complexes from a practical point of view is their ability to obtain electrons from abundant water. The utilization or imitation of the PSII functionality seems promising for all of the following: generating photoelectricity, photo-producing...

Photosynthetic response of lutein-deficient mutant lut2 of Arabidopsis thaliana to low temperature at high light

A.V. POPOVA, R. VLADKOVA, P. BORISOVA, K. GEORGIEVA, G. MIHAILOVA, V. VELIKOVA, T. TSONEV, A.G. IVANOV

Photosynthetica 2022, 60(1):110-120 | DOI: 10.32615/ps.2022.009  

Alterations in photosynthetic performance of lutein-deficient mutant lut2 and wild type (wt) of Arabidopsis thaliana were followed after treatment with low temperature and high light for 6 d. The obtained results indicated lower electrolyte leakage, lower excitation pressure, and higher actual photochemical efficiency of PSII in lut2 plants exposed to combined stress compared to wt plants. This implies that lut2 is less susceptible to the applied stress conditions. The observed lower values of quantum efficiency of nonphotochemical quenching and energy-dependent component of nonphotochemical quenching in lut2 suggest...

Sensing and classification of rice (Oryza sativa L.) drought stress levels based on chlorophyll fluorescence

Q. XIA, L.J. FU, H. TANG, L. SONG, J.L. TAN, Y. GUO

Photosynthetica 2022, 60(1):102-109 | DOI: 10.32615/ps.2022.005  

Sensing and classification of drought stress levels are very important to agricultural production. In this work, rice drought stress levels were classified based on the commonly used chlorophyll a fluorescence (ChlF) parameter (Fv/Fm), feature data (induction features), and the whole OJIP induction (induction curve) by using a Support Vector Machine (SVM). The classification accuracies were compared with those obtained by the K-Nearest Neighbors (KNN) and the Ensemble model (Ensemble) correspondingly. The results show that the SVM can be used to classify drought stress levels of rice more accurately compared to the KNN...

Variability of the photosynthetic machinery tolerance when imposed to rapidly or slowly imposed dehydration in native Mediterranean plants

E. KOUTRA, C. CHONDROGIANNIS, G. GRAMMATIKOPOULOS

Photosynthetica 2022, 60(1):88-101 | DOI: 10.32615/ps.2022.003  

Dehydration affects the photosynthetic apparatus. The impact of dehydration on photosynthesis was assessed in twelve Mediterranean species representing different growth forms. Rapid and slow dehydration experiments were conducted to (1) compare the impact of water stress among species and growth forms, (2) rank species according to their drought tolerance. Rapid dehydration reduced the electron transport up to PSI, the reduction being linearly related to leaf relative water content (RWC), except for the deciduous species. Specific energy fluxes per reaction center and maximum photochemical activity of PSII remained relatively stable until...

An open Internet of Things (IoT)-based framework for feedback controlof photosynthetic activities

S. YUAN, H. TANG, L.J. FU, J.L. TAN, G. GOVINDJEE, Y. GUO

Photosynthetica 2022, 60(1):79-87 | DOI: 10.32615/ps.2021.066  

Active control of photosynthetic activities is important in plant physiological study. Although models of plant photosynthesis have been built at different scales, they have not been fully examined for their application in plant growth control. However, we do not have an infrastructure to support such experiments since current plant growth chambers usually use fixed control protocols. In our current paper, an open IoT-based framework is proposed. This framework allows a plant scientist or agricultural engineer, through an application programming interface (API), in a desirable programming language, (1) to gather environmental data and plant...

Exposed red leaves display adaptive adjustments in chlorophyll and photosystem ratios compatible with the shade imposed by anthocyanin accumulation

K. ZELIOU, A. KYZERIDOU, Y. PETROPOULOU

Photosynthetica 2022, 60(1):70-78 | DOI: 10.32615/ps.2021.062  

Foliar anthocyanins shape a peculiar shade in a red leaf's interior leading to uneven energy distribution between the two photosystems. Accordingly, a readjustment of PSII/PSI stoichiometry could restore excitation balance. To test this hypothesis, 77 K fluorescence emission spectra of thylakoids from green and red leaves of seven species with different pigment profiles were compared. The ratio of F686/F736 served as an indication of the PSII/PSI functional ratio. To avoid possible species-dependent differences in the measured parameters, plants showing intra-individual, intra-species, or intra-leaf variation in the expression...

Intersections: photosynthesis, abiotic stress, and the plant microbiome

B. DEMMIG-ADAMS, S.K. POLUTCHKO, M.C. ZENIR, P. FOUROUNJIAN, J.J. STEWART, M. LÓPEZ-POZO, W.W. ADAMS III

Photosynthetica 2022, 60(1):59-69 | DOI: 10.32615/ps.2021.065  

Climate change impacts environmental conditions that affect photosynthesis. This review examines the effect of combinations of elevated atmospheric CO2, long photoperiods, and/or unfavorable nitrogen supply. Under moderate stress, perturbed plant source-sink ratio and redox state can be rebalanced but may result in reduced foliar protein content in C3 plants and a higher carbon-to-nitrogen ratio of plant biomass. More severe environmental conditions can trigger pronounced photosynthetic downregulation and impair growth. We comprehensively evaluate available evidence that microbial partners may be able to support plant productivity...

Light quality, oxygenic photosynthesis and more

D. LAZAR, A. STIRBET, L.O. BJÖRN, G. GOVINDJEE

Photosynthetica 2022, 60(1):25-58 | DOI: 10.32615/ps.2021.055  

Oxygenic photosynthesis takes place in thylakoid membranes (TM) of cyanobacteria, algae, and higher plants. It begins with light absorption by pigments in large (modular) assemblies of pigment-binding proteins, which then transfer excitation energy to the photosynthetic reaction centers of photosystem (PS) I and PSII. In green algae and plants, these light-harvesting protein complexes contain chlorophylls (Chls) and carotenoids (Cars). However, cyanobacteria, red algae, and glaucophytes contain, in addition, phycobiliproteins in phycobilisomes that are attached to the stromal surface of TM, and transfer excitation energy to the reaction centers via...

George C. Papageorgiou and the protective role of glycine betaine in activation and stabilization of the oxygen-evolving photosystem II complex

N. MURATA, K. STAMATAKIS

Photosynthetica 2022, 60(1):21-24 | DOI: 10.32615/ps.2021.056  

We present here our adventures in research in photosynthesis with George C. Papageorgiou (1933-2020) focusing on George's initiative in the discovery of the protective effects of glycine betaine on the oxygen-evolving photosystem II complex. We end with a brief description of research on glycine betaine-synthesizing transgenic cyanobacteria. Two of us, Norio Murata (in Japan) and Kostas Stamatakis (in Greece), and all our collaborators, have the highest respect for George, and we miss him and our intense discussions with him on various topics of photosynthesis research.

Differences in susceptibility to photoinhibition do not determinegrowth rate under moderate light in batch or turbidostat - a studywith five green algae

H. MATTILA, D. VALEV, K.B. MISHRA, V. HAVURINNE, O. VIRTANEN, M. ANTINLUOMA, E. TYYSTJÄRVI

Photosynthetica 2022, 60(1):10-20 | DOI: 10.32615/ps.2021.054  

To understand growth limitations of photosynthetic microorganisms, and to investigate whether batch growth or certain photosynthesis-related parameters predict a turbidostat (continuous growth at constant biomass concentration) growth rate, five green algal species were grown in a photobioreactor in batch and turbidostat conditions and their susceptibilities to photoinhibition of photosystem II as well as several photosynthetic parameters were measured. Growth rates during batch and turbidostat modes varied independently of each other; thus, a growth rate measured in a batch cannot be used to determine the continuous growth rate. Greatly different...

Contents of photosynthetic pigments and ratios of chlorophyll a/b and chlorophylls to carotenoids (a+b)/(x+c) in C4 plants as compared to C3 plants

H.K. LICHTENTHALER, F. BABANI

Photosynthetica 2022, 60(1):3-9 | DOI: 10.32615/ps.2021.041  

The content of chlorophylls (Chl) (a+b), total carotenoids (x+c), and the pigment ratios of Chl a/b and Chls to carotenoids (a+b)/(x+c) of green leaves of five C4 plants were determined and compared to those of C3 plants. The C4 plants were: Pacific and Chinese silvergrass (Miscanthus floridulus and Miscanthus sinensis), sugar cane (Saccharum officinarum) as well as feed and sugar maize (Zea mays). The three C3 plants were beech, ginkgo, and oak. C4 plants possess higher values for the ratio Chl a/b...

EDITORIAL

O. PRÁŠIL, RADEK KAŇA, G. GOVINDJEE

Photosynthetica 2022, 60(1):1-2 | DOI: 10.32615/ps.2022.017