PGR5/PGRL1介导的环式电子传递研究进展
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摘要
环式电子传递(CEF)是光合作用光反应中仅次于线性电子传递(LEF)的另一条重要的电子传递途径。CEF包括NAD(P)H脱氢酶复合体(NDH)和PGR5 (Proton Gradient Regulation 5)/PGRL1 (Proton Gradient Regulation Like 1)两条途径,其中PGR5/PGRL1被认为是高等植物CEF中的主要途径。本文综述了PGR5/PGRL1介导的CEF的主要作用机理,以及在逆境胁迫下该途径通过保护光系统Ⅱ和光系统Ⅰ、维持ATP/NADPH比例、调节LEF和CEF的转换来缓解光抑制中的生理作用,以期为今后的相关研究提供参考。
Cyclic electron flow(CEF) is an important electron transport pathway in light reaction of photosynthesis followed by linear electron flow(LEF). CEF includes NAD(P)H dehydrogenase complex(NDH) and PGR5(Proton Gradient Regulation 5)/PGRL1(Proton Gradient Regulation Like 1) pathways. Among them,PGR5/PGRL1-dependent CEF is considered to be the key pathway in plants. This review summarizes the mechanism of PGR5/PGRL1-dependent CEF and the physiological mechanism of PGR5/PGRL1-dependent CEF under stresses by protecting photosystem Ⅱ(PSⅡ) and PSI, maintaining ATP/NADPH ratio, and adjusting the conversion of LEF and CEF to alleviate the physiological role of photoinhibition. This review would provide a theoretical basis for future researches in the field of the mechanism of PGR5/PGRL1-dependent CEF.
Cyclic electron flow(CEF) is an important electron transport pathway in light reaction of photosynthesis followed by linear electron flow(LEF). CEF includes NAD(P)H dehydrogenase complex(NDH) and PGR5(Proton Gradient Regulation 5)/PGRL1(Proton Gradient Regulation Like 1) pathways. Among them,PGR5/PGRL1-dependent CEF is considered to be the key pathway in plants. This review summarizes the mechanism of PGR5/PGRL1-dependent CEF and the physiological mechanism of PGR5/PGRL1-dependent CEF under stresses by protecting photosystem Ⅱ(PSⅡ) and PSI, maintaining ATP/NADPH ratio, and adjusting the conversion of LEF and CEF to alleviate the physiological role of photoinhibition. This review would provide a theoretical basis for future researches in the field of the mechanism of PGR5/PGRL1-dependent CEF.
引文
Arnon DI,Allen MB,Whatley FR(1954).Photosynthesis by isolated chloroplasts.Nature,174:394-396
Arnon DI,Whatley FR,Allen MB(1958).Assimilatory power in photosynthesis.Science,127(3305):1026-1034
Asada Y,Miyake M,Miyake J,et al(1999).Photosynthetic accumulation of poly-(hydroxybutyrate)by cyanobacteria-the metabolism and potential for CO2 recycling.Int J Biol Macromol,25(1-3):37-42
Avenson TJ,Cruz JA,Kanazawa A,et al(2005).Regulating the proton budget of higher plant photosynthesis.Proc Natl Acad Sci USA,102(27):9709-9713
Bendall DS,Manasse RS(1995).Cyclic photophosphorylation and electron transport.Biochim Biophys Acta,1229(1):23-38
B?hme H,Cramer WA(1972).Localization of a site of energy coupling between plastoquinone and cytochrome f in the electron-transport chain of spinach chloroplasts.Biochemistry,11(7):1155-1160
Burrows PA,Sazanov A,Svab Z,et al(1998).Identification of a functional respiratory complex in chloroplasts through analysis of tobacco mutants containing disrupted plastid ndh genes.EMBO J,17(4):868-876
Chaux F,Johnson X,Auroy P,et al(2017).PGRL1 and LHCSR3 compensate for each other in controlling photosynthesis and avoiding Photosystem I photoinhibition during high light acclimation of Chlamydomonas cells.Mol Plant,10:216-218
Chow WS,Hope AB(2004).Electron fluxes through photosystem I in cucumber leaf discs probed by far-red light.Photosynth Res,81:77-89
DalCorso G,Pesaresi P,Masiero S,et al(2008).A complex containing PGRL1 and PGR5 is involved in the switch between linear and cyclic electron flow in Arabidopsis.Cell,132(2):273-285
Ettinger WF,Clear AM,Fanning KJ,et al(1999).Identification of a Ca2+/H+antiport in the plant chloroplast thylakoid membrane.Plant Physiol,119(4):1379-1386
Finazzi G,Rappaport F,Furia A,et al(2002).Involvement of state transitions in the switch between linear and cyclic electron flow in Chlamydomonas reinhardtii.EMBORep,3:280-285
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Friso G,Majeran W,Huang M,et al(2010).Reconstruction of metabolic pathways,protein expression,and homeostasis machineries across maize bundle sheath and mesophyll chloroplasts:large-scale quantitative proteomics using the first maize genome assembly.Plant Physiol,152:1219-1250
Gollan PJ,Lima-Melo Y,Tiwari A,et al(2017).Interaction between photosynthetic electron transport and chloroplast sinks triggers protection and signalling important for plant productivity.Philos Trans R Soc B-Biol Sci,372(1730):20160390
Govindjee(2004).Chlorophyll a fluorescence:a bit of basics and history.In:Papageorglou GC,Govindiee(eds).Chlorophyll a Fluorescence:A Signature of Photosynthesis.Dordrecht:Springer,1-42
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Hind G,Nakatani HY,Izawa S(1974).Light-dependent redistribution of ions in suspensions of chloroplast thylakoid membranes.Proc Natl Acad Sci USA,71(4):1484-1488
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Jin Y,Chen S,Fan X,et al(2017).Diuron treatment reveals the different roles of two cyclic electron transfer pathways in photosystem Ⅱ in Arabidopsis thaliana.Pestic Biochem Physiol,137:15-20
Kawashima R,Sato R,Harada K,et al(2017).Relative contributions of PGR5-and NDH-dependent photosystem Icyclic electron flow in the generation of a proton gradient in Arabidopsis chloroplasts.Planta,246:1045-1050
Kono M,Noguchi K,Terashikma I(2014).Roles of the cyclic electron flow around PSI(CEF-PSI)and O2-dependent alternative pathways in regulation of the photosynthetic electron flow in short-term fluctuating light in Arabidopsis thaliana.Plant Cell Physiol,55:990-1004
Kono M,Yamori W,Suzuki Y,et al(2017).Photoprotection of PSI by far-red light against the fluctuating light-induced photoinhibition in Arabidopsis thaliana and field-grown plants.Plant Cell Physiol,58:35-45
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Krieger A,Weis E(1993).The role of calcium in the pH-dependent control of photosystem Ⅱ.Photosynth Res,37(2):117-130
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Avenson TJ,Cruz JA,Kanazawa A,et al(2005).Regulating the proton budget of higher plant photosynthesis.Proc Natl Acad Sci USA,102(27):9709-9713
Bendall DS,Manasse RS(1995).Cyclic photophosphorylation and electron transport.Biochim Biophys Acta,1229(1):23-38
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Chaux F,Johnson X,Auroy P,et al(2017).PGRL1 and LHCSR3 compensate for each other in controlling photosynthesis and avoiding Photosystem I photoinhibition during high light acclimation of Chlamydomonas cells.Mol Plant,10:216-218
Chow WS,Hope AB(2004).Electron fluxes through photosystem I in cucumber leaf discs probed by far-red light.Photosynth Res,81:77-89
DalCorso G,Pesaresi P,Masiero S,et al(2008).A complex containing PGRL1 and PGR5 is involved in the switch between linear and cyclic electron flow in Arabidopsis.Cell,132(2):273-285
Ettinger WF,Clear AM,Fanning KJ,et al(1999).Identification of a Ca2+/H+antiport in the plant chloroplast thylakoid membrane.Plant Physiol,119(4):1379-1386
Finazzi G,Rappaport F,Furia A,et al(2002).Involvement of state transitions in the switch between linear and cyclic electron flow in Chlamydomonas reinhardtii.EMBORep,3:280-285
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Friso G,Majeran W,Huang M,et al(2010).Reconstruction of metabolic pathways,protein expression,and homeostasis machineries across maize bundle sheath and mesophyll chloroplasts:large-scale quantitative proteomics using the first maize genome assembly.Plant Physiol,152:1219-1250
Gollan PJ,Lima-Melo Y,Tiwari A,et al(2017).Interaction between photosynthetic electron transport and chloroplast sinks triggers protection and signalling important for plant productivity.Philos Trans R Soc B-Biol Sci,372(1730):20160390
Govindjee(2004).Chlorophyll a fluorescence:a bit of basics and history.In:Papageorglou GC,Govindiee(eds).Chlorophyll a Fluorescence:A Signature of Photosynthesis.Dordrecht:Springer,1-42
Hashimoto M,Endo T,Peltier G,et al(2003).A nucleus-encoded factor,CRR2,is essential for the expression of chloroplast ndhB in Arabidopsis.Plant J,36(4):541-549
Hertle AP,Blunder T,Wunder T,et al(2013).PGRL1 is the elusive ferredoxin-plastoquinone reductase in photosynthetic cyclic electron flow.Mol Cell,49(3):511-523
Hind G,Nakatani HY,Izawa S(1974).Light-dependent redistribution of ions in suspensions of chloroplast thylakoid membranes.Proc Natl Acad Sci USA,71(4):1484-1488
Huang LS,Cobessi D,Tung EY,et al(2005).Binding of the respiratory chain inhibitor antimycin to the mitochondrial bc1 complex:a new crystal structure reveals an altered intramolecular hydrogen-bonding pattern.J Mol Biol,351(3):573-597
Huang W,Yang YJ,Hu H,et al(2015).Different roles of cyclic electron flow around photosystem I under sub-saturating and saturating light intensities in tobacco leaves.Front Plant Sci,6:923
Iwai M,Takizawa K,Tokutsu R,et al(2010).Isolation of the elusive supercomplex that drives cyclic electron flow in photosynthesis.Nature,464:1210-1213
Jin Y,Chen S,Fan X,et al(2017).Diuron treatment reveals the different roles of two cyclic electron transfer pathways in photosystem Ⅱ in Arabidopsis thaliana.Pestic Biochem Physiol,137:15-20
Kawashima R,Sato R,Harada K,et al(2017).Relative contributions of PGR5-and NDH-dependent photosystem Icyclic electron flow in the generation of a proton gradient in Arabidopsis chloroplasts.Planta,246:1045-1050
Kono M,Noguchi K,Terashikma I(2014).Roles of the cyclic electron flow around PSI(CEF-PSI)and O2-dependent alternative pathways in regulation of the photosynthetic electron flow in short-term fluctuating light in Arabidopsis thaliana.Plant Cell Physiol,55:990-1004
Kono M,Yamori W,Suzuki Y,et al(2017).Photoprotection of PSI by far-red light against the fluctuating light-induced photoinhibition in Arabidopsis thaliana and field-grown plants.Plant Cell Physiol,58:35-45
Kramer DM,Avenson TJ,Edwards GE(2004).Dynamic flexibility in the light reactions of photosynthesis governed by both electron and proton transfer reactions.Trends Plant Sci,9(7):349-357
Kramer DM,Cruz JA,Kanazawa A(2003).Balancing the central roles of the thylakoid proton gradient.Trends Plant Sci,8(1):27-32
Krieger A,Weis E(1993).The role of calcium in the pH-dependent control of photosystem Ⅱ.Photosynth Res,37(2):117-130
Kukuczka B,Magneschi L,Petroutsos D,et al(2014).Proton gradient regulation5-like1-mediated cyclic electron flow is crucial for acclimation to anoxia and complementary to nonphotochemical quenching in stress adaptation.Plant Physiol,165(4):1604-1617
Labs M,Rühle T,Leister D(2016).The antimycin A-sensitive pathway of cyclic electron flow:from 1963 to 2015.Photosynth Res,129(3):231-238
Lapashina AS,Feniouk BA(2018).ADP-inhibition of H+-FOF1-ATP synthase.Biochemistry Mosc,83(10):1141-1160
Liu J,Yang YH,Hao YC,et al(2017).Effects of photosynthetic electron transport chain of maize leaves under drought stress and re-watering.Plant Physiol J,53(10):1877-1884(in Chinese with English abstract)[刘佳,杨永红,郝引川等(2017).干旱胁迫和复水处理对玉米叶片光合电子传递特性的影响.植物生理学报,53(10):1877-1884]
Lu T,Shi JW,Sun ZP,et al(2017).Response of linear and cyclic electron flux to moderate high temperature and high light stress in tomato.J Zhejiang Univ Sci B,18(7):635-648
Meng XL,Zhang LT,Zhang ZS,et al(2012).Role of mitochondrial alternative oxidase(AOX)pathway in photoprotection in Rumex K-1 leaves.Chin J Appl Ecol,23(7):1803-1808(in Chinese with English abstract)[孟祥龙,张立涛,张子山等(2012).杂交酸模叶片线粒体交替氧化酶呼吸途径在光破坏防御中的作用.应用生态学报,23(7):1803-1808]
Mosebach L,Heilmann C,Mutoh R,et al(2017).Association of ferredoxin:NADP+oxidoreductase with the photosynthetic apparatus modulates electron transfer in Chlamydomonas reinhardtii.Photosynth Res,134(3):291-306
Moss DA,Bendall DS(1984).Cyclic electron transport in chloroplasts.The Q-cycle and the site of action of antimycin.Biochim Biophys Acta,767(3):389-395
Munekage Y,Hashimoto M,Miyake C,et al(2004).Cyclic electron flow around photosystem I is essential for photosynthesis.Nature,429(3991):579-582
Munekage Y,Hojo M,Meurer J,et al(2002).PGR5 is involved in cyclic electron flow around photosystem I and is essential for photoprotection in Arabidopsis.Cell,110(3):361-371
Munekage YN,Genty B,Peitier G(2008).Effect of PGR5impairment on photosynthesis and growth in Arabidopsis thaliana.Plant Cell Physiol,49(11):1688-1698
Nandha B,Finazzi G,Joliot P,et al(2007).The role of PGR5in the redox poising of photosynthetic electron transport.Biochim Biophys Acta,1767(10):1252-1259
Nishikawa Y,Yamamoto H,Okegawa Y,et al(2012).PGR5-dependent cyclic electron transport around PSIcontributes to the redox homeostasis in chloroplasts rather than CO2 fixation and biomass production in rice.Plant Cell Physiol,53(12):2117-2126
Okegawa Y,Kagawa Y,Kobayashi Y,et al(2008).Characterization of factors affecting the activity of photosystem Icyclic electron transport in chloroplasts.Plant Cell Physiol,49(5):825-834
Okegawa Y,Kobayashi Y,Shikkanai T(2010).Physiological links among alternative electron transport pathways that reduce and oxidize plastoquinone in Arabidopsis.Plant J,63(3):458-468
Paredes M,Quiles MJ(2017).Chilling stress and hydrogen peroxide accumulation in Chrysanthemum morifolium and Spathiphyllum lanceifolium.Involvement of chlororespiration.J Plant Physiol,211:36-41
Peltier G,Tolleter D,Billon E,et al(2010).Auxiliary electron transport pathways in chloroplasts of microalgae.Photosynth Res,106:19-31
Rieske JS(1980).Inhibitors of respiration at energy-coupling site 2 of the respiratory chain.Pharmacol Ther,11(2):415-450
Rott M,Martins NF,Thiele W,et al(2011).ATP synthase repression in tobacco restricts photosynthetic electron transport,CO2 assimilation,and plant growth by overacidification of the thylakoid lumen.Plant Cell,23:304-321
Shikanai T(2007).Cyclic electron transport around photosystem I:genetic approaches.Annu Rev Plant Biol,58:199-217
Shikanai T(2016).Regulatory network of proton motive force:contribution of cyclic electron transport around photosystem I.Photosynth Res,129(3):253-2060
Sonoike K(2011).Photoinhibition of photosystem I.Physiol Plant,142:56-64
Storti M,Alboresi A,Gerotto C,et al(2018).Role of cyclic and pseudo-cyclic electron transport in response to dynamic light changes in Physcomitrella patens.Plant Cell Environ,doi:10.1111/pce.13493
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