Abstract
The high-light-induced alterations in photosynthetic performance of photosystem II (PSII) and photosystem I (PSI) as well as effectiveness of dissipation of excessive absorbed light during illumination for different periods of time at room (22 °C) and low (8–10 °C) temperature of leaves of Arabidopsis thaliana, wt and lut2, were followed with the aim of unraveling the role of lutein in the process of photoinhibition. Photosynthetic parameters of PSII and PSI were determined on whole leaves by PAM fluorometer and oxygen evolving activity—by a Clark-type electrode. In thylakoid membranes, isolated from non-illuminated and illuminated for 4.5 h leaves of wt and lut2 the photochemical activity of PSII and PSI and energy interaction between the main pigment–protein complexes was determined. Results indicate that in non-illuminated leaves of lut2 the maximum rate of oxygen evolution and energy utilization in PSII is lower, excitation pressure of PSII is higher and cyclic electron transport around PSI is faster than in wt leaves. Under high-light illumination, lut2 leaves are more sensitive in respect to PSII performance and the extent of increase of excitation pressure of PSII, ΦNO, and cyclic electron transport around PSI are higher than in wt leaves, especially when illumination is performed at low temperature. Significant part of the excessive light energy is dissipated via mechanism, not dependent on ∆pH and to functioning of xanthophyll cycle in LHCII, operating more intensively in lut2 leaves.
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Abbreviations
- BQ:
-
1,4-Benzoquinone
- CEF:
-
Cyclic electron flow around PSI
- DCMU:
-
3-(3,4-dichlorophenyl)1,1-dimethyl urea
- DCPIP:
-
2,6-dichlorophenolindophenol
- EDTA:
-
Ethylenediamine-tetraacetic acid
- ETR:
-
Electron transport rate
- F o :
-
Minimum yield of chlorophyll fluorescence in open PSII centers
- F m :
-
Maximal chlorophyll fluorescence in dark-adapted state
- \(F^{\prime}_{{\text{m}}}\) :
-
Maximal chlorophyll fluorescence in light-adapted state
- F v :
-
Variable chlorophyll fluorescence
- ΦPSII :
-
Effective quantum yield of PSII
- ΦNPQ :
-
Quantum yield of the regulated energy dissipation of PSII
- ΦNO :
-
Quantum yield of non-regulated energy dissipation of PSII
- F v/F m :
-
Maximum photochemical efficiency of PSII in the dark-adapted state
- LCP:
-
Light compensation point
- LHCII:
-
Light-harvesting chlorophyll a/b-protein complex of PSII
- LHCI:
-
Light-harvesting chlorophyll a/b-protein complex of PSI
- MES:
-
2(N-morpholino)ethanesulfonic acid
- MV:
-
Methyl viologen
- NPQ:
-
Non-photochemical quenching
- P700:
-
Reaction center chlorophyll of PSI
- P700+ :
-
Oxidized form of PSI reaction center
- PFD:
-
Photon flux density
- PQ:
-
Plastoquinone
- PSI:
-
Photosystem I
- PSII:
-
Photosystem II
- Q A, Q B :
-
Primary and secondary electron-accepting quinone in PSII
- TES:
-
N-tris[hydroxymethyl]methyl-2-aminoethanesulfonic acid
- TRICINE:
-
N-tris[hydroxymethyl]methyl glycine
References
Adams WW III, Zarter CR, Mueh KE, Amiard V, Demmig-Adams B (2008) Energy dissipation and photoinhibition: a continuum of photoprotection. In: Demmig-Adams B, Adams W, Mattoo A (eds) Photoprotection, photoinhibition, gene regulation, and environment, Springer, Dordrecht, pp 49–64
Allen JF (1995) Thylakoid protein phosphorylation, state 1- state 2 transitions, and photosystem stoichiometry adjustment: redox control at multiple levels of gene expression. Physiol Plant 93:196–205
Allen DJ, Ort DR (2001) Impacts of chilling temperatures on photosynthesis in warm-climate plants. Trends Plant Sci 6(1):36–42
Andersson J (1981) Consequence of spatial separation of photosystem I and II in thylakoid membranes from higher plants chloroplasts. FEBS Lett 124:1–10
Andrizhiyevskaya EG, Chojnicka A, Bautista JA, Diner BA, van Grondelle R, Dekker JP (2005) Origin of the F685 and F695 fluorescence in photosystem II. Photosynth Res 84:173–180
Aro E-M, Virgin I, Andersson B (1993) Photoinhibition of photosystem II. Inactivation, protein damage and turnover. Biochim Biophys Acta 1143:113–134
Baker NR, East TM, Long SP (1983) Chilling damage to photosynthesis in young Zea mays II. Photochemical function of thylakoids in vivo. J Exp Bot 34:189–197
Barenyi B, Krause GH (1985) Inhibition of photosynthetic reactions by light. A study with isolated spinach chloroplasts. Planta 163:218–226
Bassi R, Pineau B, Dainese P, Marquardt J (1993) Carotenoid binding proteins of photosystem II. Eur J Biochem 212:297–303
Bravo LA, Saavedra-Mella FA, Vera F, Guerra A, Cavieres LA, Ivanov AG, Huner NPA, Corcuera LJ (2007) Effect of cold acclimation on the photosynthetic performance of two ecotypes of Colobanthus quitensis (Kunth) Bartl. J Exp Bot 58(13):3581–3590
Brestic M, Zivcak M, Olsovska K, Shao H-B, Kalaji HM, Allakhverdiev SI (2014) Reduced glutamine synthetase activity plays a role in control of photosynthetic responses to high light in barley leaves. Plant Physiol Biochem 81:74–83
Brestic M, Zivcak M, Kunderlikova K, Sytar O, Shao H, Kalaji H, Allakhverdiev SI (2015) Low PSI content limits the photoprotection of PSI and PSII in early growth stages of chlorophyll b-deficient wheat mutant lines. Photosynth Res 125:151–166
Bruce D, Samson G, Carpenter C (1997) The origins of non-photochemical quenching of chlorophyll in photosynthesis. Direct quenching by P680(+) in photosystem II enriched membranes at low pH. Biochemistry 36:749–755
Bukhov N, Egorova E, Carpentier R (2002) Electron flow to photosystem I from stromal reductants in vivo: the size of the pool of stromal reductants controls the rate of electron donation to both rapidly and slowly reducing photosystem I units. Planta 215:812–820
Dall’Osto L, Lico C, Alric J, Giuliano G, Havaux M, Bassi R (2006) Lutein is needed for efficient chlorophyll triplet quenching in the major LHCII antenna complex of higher plants and effective photoprotection in vivo under strong light. MBC Plant Biol 6:32
Dall’Osto L, Fiore A, Cazzaniga S, Giuliano G, Bassi R (2007) Different roles of α- and β-branch xanthophylls in photosystem assembly and photoprotection. J Biol Chem 282:35056–35068
Dall’Osto L, Ünlü C, Cazzaniga S, van Amerongen H (2014) Disturbed excitation energy transfer in Arabidopsis thaliana mutants lacking antenna complexes of photosystem II. Biochim Biophys Acta 1837:1981–1988
Dekker JP, Boekema EJ (2005) Supramolecular organization of thylakoid membrane proteins in green plants. Biochim Biophys Acta 1706:12–39
Delrieu MJ (1998) Regulation of thermal dissipation of absorbed excitation energy and violaxanthin deepoxidation in the thylakoids of Lactuca sativa. Photoprotective mechanism of a population of photosystem II centers. Biochim Biophys Acta 1363:157–173
Demming-Adams B, Adams WW (1992) Photoprotection and other responses of plants to high light stress. Annu Rev Plant Physiol Plant Mol Biol 43:599–626
Derks A, Schaven K, Bruce D (2015) Diverse mechanisms for photoprotection in photosynthesis. Dynamic regulation of photosystem II excitation in response to rapid environmental change. Biochim Biophys Acta 1847:468–485
Dobrev K, Stanoeva D, Velitchkova M, Popova AV (2016) The lack of lutein accelerates the extent of light-induced bleaching of photosynthetic pigments in thylakoid membranes of Arabidopsis thaliana. Photochem Photobiol 92:436–445
Endo T, Shikanai T, Takabayashi A, Asada K, Sato F (1999) The role of chloroplastic NAD(P)H dehydrogenase in photoprotection. FEBS Lett 457:5–8
Fan D-Y, Hope AB, Jia H, Chow WS (2008) Separation of light-induced linear, cyclic and stroma-sourced electron fluxes to P700+ in cucumber leaf discs after pre-illumination at low temperature. Plant Cell Physiol 49:901–911
Frank HA, Cogdell RJ (1996) Carotenoids in photosynthesis. Photochem Photobiol 63:257–264
Genty B, Briantais J-M, Baker NR (1989) The relationship between the quantum yield of photosynthetic electron transport and quenching of chlorophyll fluorescence. Biochim Biophys Acta 990:87–92
Giersch C, Krause GH (1991) A simple model relating photoinhibitory fluorescence quenching in chloroplasts to a population of altered photosystem II reaction centers. Photosynth Res 30:115–121
Gilmore AM (1997) Mechanistic aspects of xanthophylls cycle-dependent photoprotection in higher plant chloroplasts and leaves. Physiol Plant 99:197–209
Gray GR, Savitch LV, Ivanov AG, Hüner NPA (1996) Photosystem II excitation pressure and development of resistance to photoinhibition. II. Adjustment of photosynthetic capacity in winter wheat and winter rye. Plant Physiol 110:61–71
Groce R, Weiss S, Bassi R (1999) Carotenoid-binding sites of the major light-harvesting complex II of higher plants. J Biol Chem 274:29613–29623
Hakala M, Tuominen I, Keränen M, Tyystjärvi T, Tyystjärvi E (2005) Evidence for the role of the oxygen-evolving manganese complex in photoinhibition of photosystem II. Biochim Biophys Acta 1706:68–80
Haldrup A, Jensen PE, Lunde C, Scheller HV (2001) Balance of power: a view of the mechanism of photosynthetic state transitions. Trends Plant Sci 6:301–305
Havaux M, Niyogi K (1999) The violaxanthin cycle protects plants from photooxidative damage by more than one mechanism. Proc Natl Acad Sci USA 96:8762–8767
Havaux M, DallÓsto L, Cuine S, Giuliano G, Bassi R (2004) The effect of zeaxanthin as the only xanthophyll on the structure and function of the photosynthetic apparatus in Arabidopsis thaliana. J Biol Chem 279(14):13878–13888
Hendrickson L, Furbank RT, Show WS (2004) A simple alternative approach to assessing the fate of absorbed light energy using chlorophyll fluorescence. Photosynth Res 82:73–81
Horton P, Ruban A (1992) Regulation of photosystem II. Photosynth Res 34:375–385
Horton P, Ruban AV, Walters RG (1996) Regulation of light harvesting in green plants. Annu Rev Plant Physiol Plant Mol Biol 47:655–684
Huang H-Y, Zhang Q, Zhao LP, Feng J-N, Peng C-L (2010) Does lutein plays a key role in the protection of photosynthetic apparatus in Arabidopsis under severe oxidative stress? Pak J Bot 42:2765–2774
Huang W, Yang YJ, Zhang SB (2017) Specific roles of cyclic electron flow around photosystem I in photosynthetic regulation in immature and mature leave. J Plant Physiol 209:76–83
Hundal T, Virgin I, Stryng S, Andersson B (1990) Changes of the organization of photosystem II following light-induced D1-protein degradation. Biochim Biophys Acta 1017:235–241
Huner NPA, Maxwell DP, Gray GR, Savich LV, Krol M, Ivanov AG, Falk S (1996) Sensing environmental change: PSII excitation pressure and redox signaling. Physiol Plant 98:358–364
Huner NPA, Oquist G, Sarhan F (1998) Energy balance and acclimation to light and cold. Trends Plant Sci 3:224–230
Ilioaia C, Johnson MP, Liao P-N, Pascal AA, van Grondelle R, Walla PJ, Ruban AV, Robert B (2011) Photoprotection in plants involves a change in lutein1 binding domain in the major light-harvesting complex of photosystem II. J Biol Chem 286:27247–27254
Ivanov AG, Morgan R, Gray GR, Velitchkova MY, Huner NPA (1998) Temperature/light dependent development of selective resistance of photoinhibition of photosystem I. FEBS Lett 430:288–292
Ivanov AG, Sane P, Hurry V, Krol M, Sveshnikov D, Huner NPA, Öquist G (2003) Low-temperature modulation of the redox properties of the acceptor side of photosystem II: photoprotection through reaction centre quenching of excess energy. Physiol Plant 119:376–383
Ivanov AG, Sane PV, Hurry V, Öquist G, Huner NPA (2008) Photosystem II reaction centre quenching: mechanisms and physiological role. Photosynth Res 98:565–574
Ivanov AG, Rosso D, Savitch LV, Stachula P, Rosembert M, Oquist G, Hurry V, Huner NPA (2012) Implications of alternative electron sinks in increased resistance of PSII and PSI photochemistry to high light stress in cold-acclimated Arabidopsis thaliana. Photosynth Res 113:191–206
Jahns P, Holzwarth AR (2012) The role of xanthophylls cycle and of lutein in photoprotection of photosystem II. Biochim Biophys Acta 1817:182–193
Johnson GN (2011) Physiology of PSI cyclic electron transport in higher plants. Biochem Biophys Acta 1807:384–389
Kalituho L, Rech J, Jahns P (2007) The roles of specific xanthophylls in light utilization. Planta 225:423–439
Klughammer C, Schreiber U (1991) Analysis of light-induced absorbency changes in the near-infrared spectral region. 1. Characterization of various components in isolated chloroplasts. Z Naturforsch C 46:233–244
Krause GH, Weis E (1991) Chlorophyll fluorescence and photosynthesis: the basis. Annu Rev Plant Physiol Plant Mol Biol 42:313–349
Krieger-Liszkay A (2005) Singlet oxygen production in photosynthesis. J Exp Bot 56:337–346
Laasch H (1987) Non-photochemical quenching of chlorophyll a fluorescence in isolated chloroplasts under conditions of stressed photosynthesis. Planta 171:220–225
Laemmli U (1970) Cleavage of structural proteins during the assembly of the head of bacteriophage T4. Nature 227:680–685
Laisk A, Talts E, Oja V, Eichelmann H, Peterson RB (2010) Fast cyclic electron transport around photosystem I in leaves under far-red light: a proton-uncoupled pathway? Photosynth Res 103:79–95
Lazarova D, Stanoeva D, Popova A, Vasilev D, Velitchkova M (2014) UV-B induced alteration of oxygen evolving reactions in pea thylakoid membranes as affected by scavengers of reactive oxygen species. Biol Plant 58:319–327
Lee HY, Hong YN, Chow WS (2001) Photoinactivation of photosystem II complexes and photoprotection by non-functional neighbours in Capsicum annuum L. leaves. Planta 212:332–342
Li Z, Ahn TK, Avenson TJ, Ballottari M, Cruz JA, Kramer DM, Bassi R, Fleming GR, Keasling JD, Niyogi KK (2009) Lutein accumulation in the absence of zeaxanthin restores nonphotochemical quenching in the Arabidopsis thaliana npq1 mutant. Plant Cell 21:1798–1812
Lichtenthaler HK (1987) Chlorophylls and carotenoids: pigments of photosynthetic biomembranes. Methods Enzymol 148:350–382
Lokstein H, Tian L, Polle J, DellaPenna D (2002) Xanthophyll biosynthetic mutants of Arabidopsis thaliana: altered nonphotochemical quenching of chlorophyll fluorescence is due to changes in photosystem II antenna size and stability. Biochim Biophys Acta 1553:309–319
Long SP, Humphries S, Falkowski PG (1994) Photoinhibition of photosynthesis in nature. Annu Rev Plant Physiol Plant Mol Biol 45:633–642
Matsubara S, Chow W-S (2004) Populations of photoinactivated photosystem II reaction centers characterized by chlorophyll a fluorescence lifetime in vivo. Proc Natl Acad Sci USA 101:18234–18239
Maxwell PC, Biggins J (1976) Role of cyclic electron transport in photosynthesis as measured by turnover of P700 in vivo. Biochemistry 15:3975–3981
Melis A (1985) Functional properties of photosystem IIβ in spinach chloroplasts. Biochim Biophys Acta 808:334–342
Melis A, Homann PH (1976) Heterogeneity of the photochemical centers in system II of chloroplasts. Photochem Photobiol 23:343–350
Millaleo R, Reyes-Diaz M, Alberdi M, Ivanov AG, Krol M, Hunner NPA (2013) Excess manganese differentially inhibits photosystem I versus II in Arabidopsis thaliana. J Ex Bot 64(1):343–354
Miyake C (2010) Alternative electron flows (water-water cycle and cyclic electron flow around PSI) in photosynthesis: molecular mechanisms and physiological functions. Plant Cell Physiol 51:1951–1963
Miyake C, Horiguchi S, Makino A, Shinzaki Y, Yamamoto H, Tomizawa K (2005) Effects of light intensity on cyclic electron flow around PSI and its relationship to non-photochemical quenching of Chl fluorescence in tobacco leaves. Plant Cell Physiol 46:1819–1830
Morosinotto T, Caffari S, DallOsto L, Bassi R (2003) Mechanistic aspects of the xanthophylls dynamics of higher plant thylakoids. Physiol Plant 119:347–354
Moskalenko AA, Karapetyan NV (1996) Structural role of carotenoids in photosynthetic membranes. Z Naturforsch 51c:763–771
Munekage Y, Hojo M, Meurer J, Endo T, Tasaka M, Shikanai T (2002) PGR5 is involved in cyclic electron flow around photosystem I and is essential for photoprotection in Arabidopsis. Cell 110:361–371
Munekage Y, Hashimoto M, Miyake C, Tomizawa K, Endo T, Tasaka M, Shikanai T (2004) Cyclic electron flow around photosystem I is essential for photosynthesis. Nature 429:579–582
Neale PJ, Melis A (1991) Dynamics of photosystem II heterogeneity during photoinhibition: depletion of PSIIβ from non-appressed thylakoids during strong irradiance exposure of Chlamydomonas reinhardtii. Biochim Biophys Acta 1056:195–203
Niyogi KK (1999) Photoprotection revisited: genetic and molecular approaches. Rev Plant Physiol Plant Mol Biol 50:333–359
Niyogi KK, Bjorkman O, Grossman AR (1997) The roles of specific xanthophylls in photoprotection. Proc Natl Acad Sci USA 94:14162–14167
Niyogi KK, Shih C, Chow WS, Pogson BJ, DellaPenna D, Bjoerkman O (2001) Photoprotection in a zeaxanthin- and lutein-deficient double mutant of Arabidopsis. Photosynth Res 67:139–145
Öguist G, Huner NPA (2003) Photosynthesis of overwintering evergreen plants. Annu Rev Plant Biol 54:329–355
Ohnishi N, Allakhverdiev SI, Takahashi S, Higashi S, Watanabe M, Nishiyama Y, Murata N (2005) Two-step mechanism of photodamage to photosystem II: step 1 occurs at the oxygen-evolving complex and step 2 occurs at the photochemical reaction center. Biochemistry 44:8494–8499
Ort DR (2001) When there is too much light. Plant Physiol 125:29–32
Peng CL, Gilmore AM (2003) Contrasting changes of photosystem efficiency in Arabidopsis xanthophyll mutants at room or low temperature under high irradiance stress. Photosynthetica 41(2):233–239
Peng CL, Lin Z-F, Su Y-Z, Lin G-Z, Dou H-Y, Zhao C-X (2006) The antioxidative function of lutein: electron spin resonance studies and chemical detection. Funct Plant Biol 33:839–846
Peter GF, Thornber JP (1991) Biochemical composition and organization of higher plant photosystem II light-harvesting pigment-proteins. J Biol Chem 266:16745–16754
Peterman ELG, Dukker FM, van Grondelle R, von Amerongen H (1995) Chlorophyll a and carotenoid triplet states in light-harvesting complex II of higher plants. Biophys J 69:2670–2678
Plumley FG, Schmidt DW (1987) Reconstitution of chlorophyll a/b light-harvesting complexes: xanthophyll-dependent assembly and energy transfer. Proc Natl Acad Sci USA 84:146–150
Pogson B, McDonald KA, Truong M, Britton G, DellaPenna D (1996) Arabidopsis carotenoid mutants demonstrate that lutein is not essential for photosynthesis in higher plants. Plant Cell 8:1627–1639
Popova AV, Velitchkova M, Zeinalov Y (2007) Effect of membrane fluidity on photosynthetic oxygen production reactions. Z Naturforsch 62c:253–260
Powles SB (1984) Photoinhibition of photosynthesis induced by visible light. Annu Rev Plant Physiol 35:15–44
Ravenel J, Peltier G, Havaux M (1994) The cyclic electron pathways around photosystem I in Chlamydomonas reinhardtii as determined in vivo by photoacoustic measurements of energy storage. Planta 193:251–259
Rochaix J-D (2004) Genetics of the biogenesis and dynamics of the photosynthetic machinery in eukaryotes. Plant Cell 16:1650–1660
Sane PV, Ivanov AG, Hurry V, Huner NPA, Öquist G (2003) Changes in the redox potential of primary and secondary electron accepting quinones in photosystem II confer increased resistance to photoinhibition in low-temperature-acclimated Arabidopsis. Plant Physiol 132:2144–2151
Savitch LV, Ivanov AG, Gudynaite-Savitch L, Huner NPA, Simmonds J (2011) Cold stress effects on PSI photochemistry in Zea mays: differential increase of FQR-dependent cyclic electron flow and functional implications. Plant Cell Physiol 52:1042–1054
Siefermann-Harms D (1985) Carotenoids in photosynthesis. I. Location in photosynthetic membranes and light-harvesting function. Biochim Biophys Acta 811:325–335
Sonoike K (1998) Various aspects of inhibition of photosynthesis under light/chilling stress: “photoinhibition at chilling temperatures” versus “chilling damage in the light”. J Plant Res 111:121–129
Sonoike K, Kamo M, Hihara Y, Hiyama T, Enami I (1997) The mechanism of the degradation of PsaB gene product, one of the photosynthetic reaction center subunits of photosystem I, upon photoinhibition. Photosynth Res 53:55–63
Szyszka B, Ivanov AG, Huner NPA (2007) Psychrophily is associated with differential energy partitioning, photosystem stoichiometry and polypeptide phosphorylation in Chlamidomonas taudensis. Biochim Biophys Acta 1757:789–800
Terashima I, Funayama S, Sonoike K (1994) The site of photoinhibition in leaves of Cucumis sativus L. at low temperatures is photosystem I, not photosystem II. Planta 193(2):300–306
Tjus SE, Andersson B (1993) Loss of the trans-thylakoid proton gradient is an early event during photoinhibitory illumination of chloroplast preparations. Biochim Biophys Acta 1183:315–322
Triantaphylides C, Havaux M (2009) Singlet oxygen in plants: production, detoxification and signaling. Trends Plant Sci 14:219–228
van Kooten O, Snell JFH (1990) The use of chlorophyll fluorescence nomenclature in plant stress physiology. Photosynth Res 25:147–150
Velichkova M, Popova A (2005) High light-induced changes of 77K fluorescence spectral characteristics of thylakoid membranes with modified fluidity. Bioelectrochemistry 67:81–90
Ware MA, Dall’Osto L, Ruban AV (2016) An in vivo quantitative comparison of photoprotection in Arabidopsis xanthophyll mutants. Front Plant Sci 7:841
Wei H, Yang Y-J, Zhang S-B (2017) Specific roles of cyclic electron flow around photosystem I in photosynthetic regulation in immature and mature leaves. J Plant Physiol 209:76–83
Wientjes E, van Amerongen H, Croce R (2013) LHCII is an antenna for both photosystems after long-term acclimation. Biochim Biophys Acta 1827:420–426
Yamori W, Makino A, Shikanai T (2016) A physiological role of cyclic electron transport around photosystem I in sustaining photosynthesis under fluctuating light in rice. Sci Rep 6:20147
Yruela I, Tomás R, Sanjuán ML, Torrado E, Aured M, Picorel R (1998) The configuration of β-carotene in the photosystem II reaction center. Photochem Photobiol 68:729–737
Zeinalov Y (2002) An equipment for investigations of photosynthetic oxygen production reactions. Bulg J Plant Physiol 28:57–67
Acknowledgements
This work was partially supported by Bulgarian-Swiss Research Program, Project IZEBZO-143169/1. The seeds of the wt and mutant lut2 of A. thaliana were a generous gift from Prof. R. Bassi.
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Popova, A.V., Dobrev, K., Velitchkova, M. et al. Differential temperature effects on dissipation of excess light energy and energy partitioning in lut2 mutant of Arabidopsis thaliana under photoinhibitory conditions. Photosynth Res 139, 367–385 (2019). https://doi.org/10.1007/s11120-018-0511-2
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DOI: https://doi.org/10.1007/s11120-018-0511-2