Photosynthesis: Fundamentals and advances

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Photosynthesis: Fundamentals and advances

Md. Alamgir Hossain, Mohammed Arif Sadik Polash, Md. Masudul Karim, Sadiya Arefin Juthee, Md. Arif Sakil
Int. J. Biosci.21( 2), 443-463, August 2022.
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Abstract

Photosynthesis is one of the most incomparable and meticulous metabolic processes that maximize the use of available light, carbon and nitrogen and minimizes the destructive effects of surplus light. Indeed, photosynthesis comprises of two major reactions that occur in separate parts in the chloroplast. The light reactions take place in the thylakoid membrane which generates ATP and NADPH while dark reactions (so called Calvin–Benson cycle) exploit these ATP and NADPH to reduce carbon CO2 (carbon-di-oxide) to carbohydrates (CH2O) in the stroma of chloroplast. In plants various carbon fixation mechanisms are evolved naturally such as, less efficient C3 carbon fixation having photorespiration, more efficient C4 carbon fixation having cellular CO2 pumping system for avoiding photorespiration and CAM (Crassulacean acid metabolism) carbon fixation for escaping transpiration during day. Besides plant proceeds different alternative sinks for carbon fixation under surplus light. Chlorophyll fluorescence is one of the most influential and advanced technique for studying photosystems health but on the other hand photo inhibition and ROS (reactive oxygen species) generation are unfortunate for photosystems during various stresses. However, photo inhibition and ROS generation are obligatory during stresses whereas chloroplastic antioxidants are accountable for ROS regulation in plant cells. These insight between fundamental and advance information on photosynthesis assist to switch less efficient C3 rice to highly efficient C4 rice development to feed the ever-increasing population in the globe. Therefore, this article reviews fundamental aspects of photosynthetic machineries, underlying physiological, biochemical and molecular mechanisms and highlighted the modern scientific achievements on C4 rice development.

VIEWS 48

Adams WW III, Demmig-Adams B. 2004. Chlorophyll fluorescence as a tool to monitor plant response to the environment. In: Papageorgiou GC and Govindjee (eds) Chlorophyll a Fluorescence: A Probe of Photosynthesis. Springer, Dordrecht 583-604.

Adams WWIII, Demmig-Adams B, Winter K, Schreiber U. 1990. The ratio of variable to maximum chlorophyll fluorescence from photosystem II, measured in leaves at ambient temperature and at 77K, as an indicator of the photon yield of photosynthesis. Planta 180, 166-174.

Adams WWIII, Zarter CR, Mueh KE, Amiard V, Demmig-Adams B. 2005. Energy dissipation and photoinhibition: acontinuum of photoprotection. In: Demmig-Adams, B., Adams, W. W. III. and Mattoo, A. K. (eds) Photoprotection, Photoinhibition, Gene Regulation, and Environment. Springer, Dordrecht 49-64.

Allen JF, Forsberg J. 2001. Molecular recognition in thylakoid structure and function. Trends in Plant Science 6, 317-326.

Alscher RG. 1989. Biosynthesis and antioxidant function of glutathione in plants. Physiologia Plantarum 77, 457-64.

Anderson JM, Park YI, Chow WS. 1997. Photoinactivation and photoprotection of Photosystem II in nature. Physiologia Plantarum 100, 214-223.

Arnon DI. 1956. Phosphorus metabolism and photosynthesis. Annual Review of Plant Physiology 7(1), 325-356.

Aro EM, mcCaffery S, Anderson JM. 1993. Photoinhibition and D1protein degradation in peas acclimated to different growth irradiances. Plant Physiology 103, 835-843.

Aro EM, Virgin I, Andersson B. 1993. Photoinhibition of photosystem II, Inactivation, protein damage and turnover. Biochimica et Biophysica Acta (BBA) Bioenergetics 1143(2), 113-134.

Asada K. 1999. The water-water cycle in chloroplasts: scavenging of active oxygens and dissipation of excess photons. Annual Review of Plant Physiology and Plant Molecular Biology 50, 601-639.

Asada K. 2000. The water–water cycle as alternative photon and electron sinks. Philosophical Transactions of the Royal Society B 355, 1419-1431.

Asada K. 2006. Production and scavenging of reactive oxygen species in chloroplasts and their functions. Plant Physiology 141, 391-396.

Badger MR, Price GD. 2003. CO2 concentrating mechanisms in cyanobacteria: molecular components, their diversity and evolution. Journal of Experimental Botany 54(383), 609-622.

Bailey S, Grossman A. 2008. Photo protection in cyanobacteria: regulation of light harvesting. Photochemistry and Photobiology 84(6), 1410-1420.

Baker NR. 1996. Photo inhibition of Photosynthesis. In: Jennings, R. C., Zucchelli, G., Ghetti, F., and Colombetti, G. (eds) Light as an Energy Source and Information Carrier in Plant Physiology. NATO ASI Series (Series A: Life Sciences), vol 287. Springer, Boston, MA. Baker NR. 2008. Chlorophyll fluorescence: a probe of photosynthesis in vivo. Annual Review of Plant Biology 59, 89-113.

Barber J, Nield N, Morris EP, Hankamer B. 1999. Subunit positioning in photosystem II revisited. Trends in Biochemical Sciences 24, 43-45.

Baroli I, Niyogi KK. 2000. Molecular genetics andxanthophyll-dependent photo protection in green algae andplants. Philosophical Transactions of the Royal Society B 355, 1385-1394.

Bartley GE, Scolnik PA. 1995. Plant carotenoids: pigments for photo protection, visual attraction, and human health. Plant Cell 7, 1027-1038.

Bassham JA, Calvin M, Benson AA. 1950. The Path of Carbon in Photosynthesis: The Role of Malic Acid, University of California Radiation Laboratory.

Bastide B, Sipes D, Hann J, Ting IP. 1993. Effect of severe water stress on aspects of crassulacean acid metabolism in Xerosicyos. Plant Physiology 103(4), 1089-1096.

Bellasio C, Farquhar GD. 2019. A leaf‐level biochemical model simulating the introduction of C2 and C4 photosynthesis in C3 rice: gains, losses and metabolite fluxes. New Phytologist 223(1), 150-166.

Bellasio C, Lundgren MR. 2016. Anatomical constraints to C4 evolution: light harvesting capacity in the bundle sheath. New Phytologist 212(2), 485-496.

Bellasio C. 2017. A generalized stoichiometric model of C3, C2, C2+ C4, and C4 photosynthetic metabolism. Journal of Experimental Botany 68(2), 269-282.

Bilger W, Bjorkman O. 1990. Role of the xanthophyll cycle in photoprotection elucidated by measurements of light-induced absorbance changes, fluorescence and photosynthesis in leaves of Hedera canariensis. Photosynthesis Research 25, 173-18.

Bjorkman O, Demmig B. 1987. Photon yield of O2 evolution and chlorophyll fluorescence characteristics at 77K among vascular plants of diverse origin. Planta 170, 489-504.

Blankenship RE, Hartman H. 1998. The origin and evolution of oxygenic photosynthesis. Trends in Biochemical Sciences 23, 94-97.

Bonner W, Bonner J. 1948. The role of carbon dioxide in acid formation by succulent plants. American Journal of Botany 113-117.

Bussotti F, Gerosa G, Digrado A,  Pollastrini  M. 2020. Selection of chlorophyll fluorescence parameters as indicators of photosynthetic efficiency in large scale plant ecological studies. Ecological Indicators 108, 105686.

Callahan FE, Becker DW, Cheniae GM. 1986. Studies on the photoactivation of the water-oxidizing enzyme: II. Characterization of weak light photoinhibition of PSII and its light-induced recovery. Plant physiology 82(1), 261-269.

Calvin M, Benson AA. 1948. The Path of Carbon in Photosynthesis. Washington, DC: US.

Calvin M. 1989. Forty years of photosynthesis and related activities. Photosynthesis Researc 21(1), 3-16.

Campbell NA, Williamson B, Heyden RJ. 2006. Biology: exploring life. Recording for the Blind and Dyslexic.

Cazzaniga S, Bressan M, Carbonera D, Agostini A, Dall’Osto L. 2016. Differential roles of carotenes and xanthophylls in photosystem I photoprotection. Biochemistry 55, 3636-3649.

Chen S, Qu N, Cao S, Hermann B, Chen S, Tian W, Chu C. 2001. Expression analysis of gdcs P promoter from C3-C4 intermediate plant Flaveria anomala in transgenic rice Chinese Science Bulletin 46(19), 1635-1638.

Chen Z, Spreitzer RJ. 1992. How various factors influence the CO2/O2 specificity of ribulose-1,5-bisphosphate carboxylase/oxygenase. Photosynthesis Research 31, 157-164.

Chitnis PR. 2001. Photosystem I: Function and physiology. Annual Review of Plant Physiology and Plant Molecular Biology 52, 593-626.

Chu C, Dai Z, Ku MS, Edwards GE. 1990. Induction of Crassulacean acid metabolism in the facultative halophyte Mesembryanthemum crystallinum by abscisic acid. Plant Physiology 93(3), 1253-1260.

Clarke JE, Johnson GN. 2001. In vivo temperature dependence of cyclic and pseudocyclic electron transport in barley. Planta 212, 808-816.

Conklin PL, Saracco SA, Norris SR, Last RL. 2000. Identifi-cation of ascorbic acid-deficient Arabidopsis thaliana mutants. Genetics 154, 847-856.

Cushman JC. 2001. Crassulacean acid metabolism. A plastic photosynthetic adaptation to arid environments. Plant physiology 127(4), 1439-1448.

Danila FR, Quick WP, White RG, Kelly S, von Caemmerer S, Furbank RT. 2018. Multiple mechanisms for enhanced plasmodesmata density in disparate subtypes of C4 grasses. Journal of Experimental Botany 69(5), 1135-1145.

Demmig-Adams B, Adams WW, Barker DH, Logan BA, Bowling BA., Verhoeven AS. 1996. Using chlorophyllfluorescence to assess the fraction of absorbed light allocated to thermal dissipation of excess excitation. Physiologia Plantarum 98, 253-264.

Demmig-Adams B, Moeller DL, Logan BA, Adams WWIII. 1998. Positive correlation between levels of retained zeaxanthin+antheraxanthin and degree of photoinhibition in shade leaves of Schefflera arboricola (Hayata) Merrill. Planta 205, 367-374.

Earl HJ, Ennahli S. 2004. Estimating photosynthetic electron transport via chlorophyll fluorometry without Photosystem II light saturation. Photosynthesis Research 82(2), 177-186.

Ebbighausen H, Hatch MD, Lilley RMcC, Kro ̈mer S, Stitt M, Heldt HW. 1987. On the function of malate–oxaloacetate shuttles in a plant cell. InPlant Mitochondria: Structural, Functional and Physiological Aspects. Edited by Moore AL, Beechey RB. Plenum Press, New York 171-180.

Ehleringer JR, Sage RF, Flanagan LB, Pearcy RW. 1991. Climate change and the evolution of C4 photosynthesis. Trends in Ecology and Evolution 6(3), 95-99.

Eisenhut M, Ruth W, Haimovich M, Bauwe H, Kaplan A, Hagemann M. 2008. The photorespiratory glycolate metabolism is essential for cyanobacteria and might have been conveyed endosymbiontically to plants. Proceedings of the National Academy of Sciences 105(44), 17199-17204.

El-Sharkawy M, Hesketh J. 1965. Photosynthesis among Species in Relation to Characteristics of Leaf Anatomy and CO2 Diffusion Resistance 1. Crop Science 5(6), 517-521.

Ermakova M, Danila FR, Furbank RT, von Caemmerer S. 2020. On the road to C4 rice: advances and perspectives. The Plant Journal 101(4), 940-950.

Evans JR, Vogelmann TC, von Caemmerer S. 2008. Balancing light capture with distributed metabolic demand during C4 photosynthesis. In Charting new pathways to C4 rice 127-143.

Fajer J, Davis MS, Brune DC, Spaulding LD, Borg DC, Forman A. 1977. Chlorophyll radicals and primary events. In: Olson, J. M., and Hind, G. (eds) Brook haven symposia in biology 28, 74-104.

Fioretto A, Alfani A. 1988. Anatomy of succulence and CAM in 15 species of Senecio. Botanical Gazette 149(2), 142-152.

Flores FG. 2008. The cyanobacteria: molecular biology, genomics, and evolution, Horizon Scientific Press.

Forseth I. 2010. The ecology of photosynthetic pathways. Nature Education Knowledge 3(10), 4.

Foyer CH, Gomez LD, van Heerden PDR. 2005. Glutathione.In: Smirnoff N (ed) Antioxidants and Reactive Species in Plants. Blackwell Publishing, London.

Foyer CH, Trebst A, Noctor G. 2008. Signaling and Integration of Defense Functions ofTocopherol, Ascorbate and Glutathione. In: Demmig-Adams, B., Adams, W. W. III., and Mattoo, A. K. (eds) Photoprotection, Photoinhibition, Gene Regulation, and Environment. Springer, Dordrecht 241-268.

Frank HA, CogdeIl RJ. 1996. Carotenoids in Photosynthesis. Photochemistry and Photobiology 63(3), 257-264.

Frank HA, Cogdell RJ. 1993. The photochemistry and function of carotenoids in photosynthesis. In: Young, A., and Britton, G. (eds) Carotenoids in Photosynthesis. Champ-man and Hall, London 252-326.

Fridlyand LE, Backhausen JE, Scheibe R. 1998. Flux control of the malate valve in leaf cells. Archives of Biochemistry and Biophysics 349, 290-298.

Fryer MJ, Andrews JR, Oxborough K, Blowers DA, Baker NR. 1998. Relationship between CO2 assimilation, photosynthetic electron transport, and active O2 metabolism in leaves of maize in the field during periods of low temperature. Plant Physiology 116, 571-580.

Fryer MJ. 1992. The antioxidant effects of thylakoid vitamin E (α-tocoherol). Plant Cell and Environment 15, 381-392.

Fukayama H, Tsuchida H, Agarie S, Nomura M, Onodera H, Ono K, Lee BH, Hirose S, Toki S, Ku MSB, Makino A, Matsuoka M, Miyao M. 2001. Significant accumulation of C4-specific Pyruvate, orthophosphate dikinase in a C3 plant, rice. Plant Physiology 127, 1136-1146.

Furbank RT, von Caemmerer S, Sheehy J, Edwards G. 2009. C4 rice: a challenge for plant phenomics. Functional Plant Biology 36(11), 845-856.

Galvão VC, Fankhauser C. 2015. Sensing the light environment in plants: photoreceptors and early signaling steps. Current Opinion in Neurobiology 34, 46-53.

Gest H. 2002. History of the word photosynthesis and evolution of its definition. Photosynthesis Research 73(1), 7-10.

Green BR, Durnford DG. 1996. The chlorophyll-carotenoid proteins of oxygenic photosynthesis. Annual Review of Plant Physiology and Plant Molecular Biology 47, 685-714.

Griffiths H. 2006. Designs on Rubisco. Nature 441(7096), 940-941.

Grossman AR, Bhaya D, Apt KE, Kehoe DM. 1995. Light-harvesting complexes in oxygenic photosynthesis: Diversity, control, and evolution. Annual Review of genetics 29, 231-288.

Guerreromg, Vega JM, Losada M. 1981. The assimilatory nitrate-reducing system and its regulation. Annual Review of Plant Physiology 32, 169-202.

Guidi L, Lo Piccolo E, Landi M. 2019. Chlorophyll Fluorescence, Photoinhibition and Abiotic Stress: Does it Make Any Difference the Factto Be a C3 or C4 Species? Frontiers in Plant Science 10, 174.

Guralnick LJ, Jackson MD. 2001. The occurrence and phylogenetics of crassulacean acid metabolism in the Portulacaceae. International Journal of Plant Science 162(2), 257-262.

Gururani MA, Venkatesh J, PhanTra LS. 2015. Regulation of Photosynthesis during Abiotic Stress-Induced Photoinhibition. Molecular Plant 8(9), 1304-1320.

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. Biochimica et Biophysica Acta (BBA)-Bioenergetics 1706(1-2), 68-80.

Haldrup A, Jensen PE, Lunde C, Scheller HV. 2001. Balance of power: A view of the mechanism of photosynthetic state transitions Trends in Plant Science 6, 301-305.

Hamlyn G. Jones. 1992. Plants and microclimate: a quantitative approach to environmental plant physiology, Cambridge University Press.

Haraux F, De Kouchkovsky Y. 1998. Energy coupling and ATP synthase. Photosynthesis Research 57, 231-251.

Häusler RE, Hirsch HJ, Kreuzaler F, Peterhänsel C. 2002. Overexpression of C4‐cycle enzymes in transgenic C3 plants: a biotechnological approach to improve C3‐photosynthesis. Journal of Experimental Botany 53(369), 591-607.

Heineke D, Riens B, Grosse H, Hoferichter P, Peter U, Flügge UI, Heldt HW. 1991. Redox transfer across the inner chloroplast envelope membrane. Plant Physiology 95, 1131-1137.

Hermida-Carrera C, Kapralov MV, Galmés J. 2016. Rubisco catalytic properties and temperature response in crops. Plant Physiology 171(4), 2549-2561.

Hibberd JM, Sheehy JE, Langdale JA. 2008. Using C4 photosynthesis to increase the yield of rice-rationale and feasibility. Current Opinion in Plant Biology 11(2), 228-231.

Hill R. 1939. Oxygen produced by isolated chloroplasts. Proceedings of the Royal Society of London Series B-Biological Sciences 127(847), 192-210.

Hoganson CW, Babcock GT. 1997. A metalloradical mechanism for the generation of oxygen from water in photosynthesis. Science 277, 1953-1956.

Horton P, Ruban A. 2005. Molecular design of the photosystem II light-harvesting antenna: photosynthesis and photoprotection. Journal of Experimental Botany 56(411), 365-373.

Hossain MA, Uddin MK, Ismail MR, Ashrafuzzaman M. 2012. Responses of glutamine synthetase-glutamate synthase cycle enzymes in tomato leaves under salinity stress. International Journal of Agriculture and Biology 14(4).

Hudson D, Guevara DR, Hand AJ, Xu Z, Hao L, Chen X, Zhu T, Bi YM, Rothstein SJ. 2013. Rice cytokinin GATA transcription Factor1 regulates chloroplast development and plant architecture. Plant Physiology 162(1), 132-144.

Hultine, KR, Cushman JC, Williams DG. 2019. New perspectives on crassulacean acid metabolism biology. Journal of Experimental Botany 70(22), 6489-6493.

Igamberdiev AU. 2015. Control of Rubisco function via homeostatic equilibration of CO2 supply. Frontiers in Plant Science 6,106.

Inoue K, Sakurai H, Hiyama T. 1986. Photoinactivation sites of photosystem I in isolated chloroplasts. Plant Cell Physiology 27, 961-968.

Jalali-e-Emam SMS, Alizadeh B, Zaefizadeh M, Zakarya RA, Khayatnezhad M. 2011. Superoxide dismutase (SOD) activity in NaCl stress in salt-sensitive and salt-tolerance genotypes of Colza (Brassica napus L.). Middle-East Journal of Scientific Research 7, 7-11.

Jegerschoeld C, Virgin I, Styring S. 1990. Light-dependent degradation of the D1 protein in photosystem II is accelerated after inhibition of the water splitting reaction. Biochemistry 29(26), 6179-6186.

Johnson GN, Young AJ, Scholes JD, Horton P. 1993. The dissipation of excess excitation energy in British plant species. Plant Cell and Environment    16, 673-679.

Johnson MP, Goral TK, Duffy CD, Brain AP, Mullineaux CW, Ruban AV. 2011. Photoprotective energy dissipation involves the reorganization of photosystem II light-harvesting complexes in the grana membranes of spinach chloroplasts. The Plant Cell 23(4), 1468-1479.

Johnson MP. 2017. Correction: Photosynthesis. Essays in Biochemistry 61(4), 429-429.

Joliot P, Barbieri G, Chabaud R. 1969. A new model of photochemical centers in system-2. Photochemistry and Photobiology 10, 309-329.

Jones HG, Hillis RM, Gordon SL, Brennan RM. 2013. An approach to the determination of winter chill requirements for different Ribes cultivars. Plant Biology 15, 18-27.

Joyard J, Block MA, Douce R. 1991. Molecular aspects of plastid envelope biochemistry. European Journal of Biochemistry 199(3), 489-509.

Jung HS, Niyogi KK. 2008. Molecular Analysis of Photo protection of Photosynthesis. In: Demmig-Adams, B., Adams, W. W. III, and Mattoo, A. K. (eds), Photo protection, Photo inhibition, Gene Regulation, and Environment. Springer, Dordrecht 127-143.

Jung J, Kim HS. 1990. The chromophores as endogenous sensitizers involved in the photogeneration of singlet oxygen in spinach thylakoids. Photochemistry and Photobiology 52(5), 1003-1009.

Karplus PA, Daniels MJ, Herriott JR. 1991. Atomic structure of ferredoxin-NADP+ reductase: Prototype for a structurally novel flavor enzyme family. Science 251, 60-66.

Kebeish R, Niessen M, Thiruveedhi K, Bari R, Hirsch H J, Rosenkranz R, Stabler N, Schonfeld B, Kreuzaler F, Peterhansel C. 2007. Chloroplastic photo respiratory bypass increases photosynthesis and biomass production in Arabidopsis thaliana. Nature Biotechnology 25, 593-599.

Keeley JE, Rundel PW. 2003. Evolution of CAM and C4 carbon-concentrating mechanisms. International Journal of Plant Science 164(S3), S55-S77.

Keeley JE. 1998. CAM photosynthesis in submerged aquatic plants. Botanical Review 64(2), 121-175.

Kellogg EA. 2013. C4 photosynthesis. Current Biology 23(14), R594-R599.

Kitajima M, Butler WL. 1975. Quenching of chlorophyll fluorescence and primary photochemistry in chloroplasts by dibromothymoquinone. Biochimica et Biophysica Acta 376, 105-115.

Kok B, Forbush B, mcGloin M. 1970. Cooperation of charges in photosynthetic O2 evolution-I. A linear four step mechanism. Photochemistry and Photobiology 11, 457-475.

Kok B. 1956. On the inhibition of photosynthesis by intense light. Biochimica et Biophysica Acta 21(2), 234-244.

Krause GH, Weis E. 1991. Chlorophyll fluorescence and photosynthesis: The basics. Annual Review of Plant Physiology and Plant Molecular Biology 42, 313-350.

Krieger-Liszkay A, Fufezan C, Trebst A. 2008. Singlet oxygen production in photosystem II and related protection mechanism Photosynthesis Research 98(1), 551-564.

Krieger-Liszkay A. 2005. Singlet oxygen production in photosynthesis. Journal of Experimental Botany 56(411), 337-346.

Ku MSB, Agarie S, Nomura M, Fukayama H, Tsuchida H, Ono K, Hirose S, Toki S, Miyao M, Matsuoka M. 1999. High-level expression of maize phosphoenol pyruvate carboxylase in transgenic rice plants. Nature Biotechnology 17, 76-80.

Leegood RC. 2007. A welcome diversion from photorespiration. Nature Biotechnology 25(5), 539-540.

Lin H, Arrivault S, Coe RA, Karki S, Covshoff S, Bagunu E, Lunn JE, Stitt M, Furbank RT, Hibberd JM, Quick WP. 2020. A Partial C4 Photosynthetic Biochemical Pathway in Rice. Frontiers in Plant Science 11, 564463.

Lüttge U. 2004. Ecophysiology of crassulacean acid metabolism (CAM). Annals of Botany 93(6), 629-652.

Lyu MJA, Wang Y, Jiang J, Liu X, Chen G, Zhu XG. 2020. What Matters for C4 Transporters: Evolutionary Changes of Phosphoenol pyruvate Transporter for C4 Photosynthesis. Frontiers in Plant Science 11, 935.

Martin SL, Davis R, Prottim P, Linm TC, Lin SH, Martin CE. 2005. The occurrence of crassulacean acid metabolism in epiphytic ferns, with an emphasis on the Vittariaceae. International Journal of Plant Science 166(4), 623-630.

Matsuoka M, Furbank RT, Fukayama H, Miyao M. 2001. Molecular engineering of C4 photosynthesis. Annual Review of Plant Biology 52(1), 297-314.

Matsuoka M, Kyozuka J, Shimamoto K, Kano-Murakami Y. 1994. The promoters of two carboxylases in a C4 plant (maize) direct cell-specific, light-regulated expression in a C3 plant (rice). The Plant Journal 6, 311-319.

Maxwell K, Johnson GN. 2000. Chlorophyll fluorescence-a practical guide. Journal of Experimental Botany 51(345), 659-668

Mehler AH. 1951. Studies on reactions of illuminated chloroplasts: I. Mechanism of the reduction of oxygen and other hill reagents. Archives of Biochemistry 33(1), 65-77.

Miflin BJ, Lea PJ. 1976. The Pathway of nitrogen assimilation in plants. Phytochemistry 152(1), 873-885.

Mitchell P. 1966. Chemiosmotic coupling in oxidative and photosynthetic phosphorylation. Biological reviews of the Cambridge Philosophical Society 41, 445-501.

Miyao M, Masumoto C, Miyazawa SI, Fukayama H. 2011. Lessons from engineering a single-celled C4photosynthetic pathway into rice. Journal of Experimental Botany 62, 3021-3029.

Monson RK, Rawsthorne S. 2000. CO2 assimilation in C3-C4 intermediate plants. In: Leegood, R. C., Sharkey, T. D., and von Caem merer, S., (eds) Photosynthesis: physiology and metabolism. Kluwer Academic Publishers, Dordrecht, The Netherlands 533-550.

Müller P, Li XP, Niyogi KK. 2001. Non-photochemical quenching. A response to excess light energy. Plant physiology 125(4), 1558-1566.

Muller-Moule P, Conklin PL, Niyogi KK. 2002. Ascorbate deficiency can limit violaxanthin de-epoxidase activity in vivo. Plant Physiology 128, 970-977.

Mullineaux CW. 1999. The thylakoid membranes of cyanobacteria: structure, dynamics and function. Functional Plant Biology 26(7), 671-677.

Munekage YN, Taniguchi YY. 2016. Promotion of cyclic electron transport around photosystem I with the development of C4 photosynthesis. Plant Cell Physiology 57(5), 897-903.

Munn ́e-Bosch S, Alegre L. 2002. The function of tocopherols and tocotrienols in plants. Critical Review in Plant Science 21, 31-57.

Murata N, Allakhverdiev SI, Nishiyama Y. 2012. The mechanism of photoinhibition in vivo: Re-evaluation of the roles of catalase, α-tocopherol, non-photochemical quenching, and electron transport. Biochimica et Biophysica Acta (BBA) -Bioenergetics 1817(8), 1127-1133.

Murata N, Takahashi S, Nishiyama Y, Allakhverdiev SI. 2007. Photoinhibition of photosystem II under environmental stress. Biochimica et Biophysica Acta (BBA) -Bioenergetics 1(767), 414-421.

Murchie EH, Lawson T. 2013. Chlorophyll fluorescence analysis: a guide to good practice and understanding some new applications. Journal of Experimental Botany 64(13), 3983-3998.

Nathan N, Wolfgang J. 2015. Structure and energy transfer in photosystems of oxygenic photosynthesis. Annual Review of Biochemistry 84, 659-683.

Niyogi KK, Grossman AR, Bjorkman O. 1998. Arabidopsis mutants define a central role for the xanthophyll cycle in theregulation of photosynthetic energy conversion. The Plant Cell 10, 1121-1134.

Niyogi KK. 1999. Photoprotection revisited: genetic and molecular approaches. Annual Review of Plant Physiology and Plant Molecular Biology 50, 333-359

Niyogi KK. 2000. Safety valves for photosynthesis Current Opinion in Plant Biology 3, 455-460.

Noctor G, Foyer CH. 1998. Ascorbate and glutathione: keeping active oxygen under control. Annual Review of Plant Physiology and Plant Molecular Biology 49, 249-279.

Nomura M, Higuchi T, Ishida Y, Ohta S, Komari T, Imaizumi N, Miyao-Tokutomi M, Matsuoka M, Tajima S. 2005. Differential expression pattern of C4 bundle sheath expression genes in rice, a C3 plant. Plant Cell Physiology 46(5), 754-761.

Ohnishi N, Allakhverdiev SI, Takahashi S, Higashi S, Watanabe M, Nishiyama Y, Murata N. 2005. Two-step mechanism of photo damage to photosystem II: step 1 occurs at the oxygen-evolving complex and step 2 occurs at the photochemical reaction center. Biochemistry 44(23), 8494-8499.

Onge KRS. 2018. Photosynthesis: Light Reactions, eLS 1-8.

Ort DR. 2001. When there is too much light. Plant Physiology 125, 29-32.

Pfündel E, Bilger W. 1994. Regulation and the possible function of the violaxanthin cycle. Photosynthesis Research 42, 89-1.

Polash MAS, Sakil MA, Hossain MA. 2019. Plants responses and their physiological and biochemical defense mechanisms against salinity: A review. Tropical Plant Research 6, 250-274.

Polle A. 2001. Dissecting the superoxide-dismutase-ascorbate-glutathione pathway in chloroplasts by metabolic modeling. Computer simulations as a step towards flux analysis. Plant Physiology 126, 445-462.

Pushkar Y, Yano J, Sauer K, Boussac A, Yachandra VK. 2008. Structural changes in the Mn4Ca cluster and the mechanism of photosynthetic water splitting. Proceedings of the National Academy of Sciences 105(6), 1879-1884.

Rabinowitch EI. 1956. Photosynthesis and related processes. Vol. II. Part 2. Photosynthesis and related processes Vol. II. Part 2.

Rachmilevitch S, Cousins AB, Bloom AJ. 2004. Nitrate assimilation in plant shoots depends on photorespiration. Proceedings of the National Academy of Sciences 101(31), 11506-11510.

Ranson SL, Thomas M. 1960. Crassulacean acid metabolism, Annual Review of Plant Physiology 11(1), 81-110.

Raven PH, Evert RF, Eichhorn SE. 2005. Biology of Plants, WH Freeman and Company Publishers. New York 944.

Reeves G, Grangé-Guermente MJ, Hibberd JM. 2017. Regulatory gateways for cell-specific gene expression in C4 leaves with Kranz anatomy. Journal of Experimental Botany 68(2), 107-116.

Reyes-Prieto A, Weber AP, Bhattacharya D. 2007. The origin and establishment of the plastid in algae and plants. Annual Review of Genetics 41, 147-168.

Rich PR, Bendall DS. 1981. Electron and proton transfer in the plastoquinol-plastocyanin oxidoreductase. In Vectorial Reactions in Electron and Ion Transport in Mitochondria and Bacteria, Elsevier/North-Holland Biomedical Press, The Netherlands 187-190.

Rich PR. 1988. A critical examination of the supposed variable proton stoichiometry of the chloroplast cytochrome b/f complex. Biochimica et Biophysica Acta (BBA) – Bioenergetics 932, 33-42.

Rizal G, Karki S, Thakur V, Chatterjee J, Coe RA, Wanchana S, Quick WP. 2012. Towards a C4 rice. Asian Journal of Cell Biology 7, 13-31.

Robinson JM. 1988. Does O2 photo reduction occur within chloroplasts in vivo? Physiologia Plantarum 72, 666-680.

Robinson JM. 1990. Nitrite photo reduction in vivo is inhibited by oxygen. Plant Physiology 92, 862-865.

Rosenqvist E, van Kooten O. 2003. Chlorophyll fluorescence: a general description and nomenclature. In Practical applications of chlorophyll fluorescence in plant biology. Springer, Boston MA 31-77.

Rutherford AW, Osyczka A, Rappaport F. 2012. Back-reactions, short-circuits, leaks and other energy wasteful reactions in biological electron transfer: redox tuning to survive life in O2. FEBS Letters 586, 603-616.

Sacksteder CA, Kanazawa A, Jacoby ME, Kramer DM. 2000. The proton to electron stoichiometry of steady-state photosynthesis in living plants: a proton-pumping Q cycle is continuously engaged. Proceedings of the National Academy of Sciences of the United States of America 97, 14283-14288.

Sage RF. 2004. The evolution of C4 photosynthesis. New Phytologist 161(2), 341-370.

Sage TL, Sage RF. 2009. The functional anatomy of rice leaves: implications for refixation of photorespiratory CO2 and efforts to engineer C4 photosynthesis into rice. Plant Cell Physiology 50(4), 756-72.

Satoh K. 1970. Mechanism of photo inactivation in photosynthetic systems II. The occurrence and properties of two different types of photo inactivation. Plant Cell Physiology 11, 29-38.

Sauer K, Yachandra VK. 2004. The water-oxidation complex in photosynthesis. Biochimica et Biophysica Acta (BBA) – Bioenergetics 1665, 140-146

Scheibe R, Backhausen JE, Emmerlich V, Holtgrefe S. 2005. Strategies to maitain redox homeostasis during photo-synthesis under changing conditions. Journal of Experimental Botany 56, 1481-1489.

Schluter U, Weber APM. 2020. Regulation and evolution of C4 photosynthesis. Annual Review of Plant Biology 71(16), 1-33.

Sedelnikova OV, Hughes TE, Langdale JA. 2018. Understanding the Genetic Basis of C4 Kranz Anatomy with a View to Engineering C3 Crops. Annual Review of Genetics 52(1), 249-270.

Selinski J, Scheibe R. 2019. Malate valves: old shuttles with new perspectives. Plant Biology 21, 21-30.

Sharkey TD. 1988. Estimating the rate of photorespiration in leaves. Physiologia Plantarum 73(1), 147-152.

Shimakawa G, Miyake C. 2018. Oxidation of P700 Ensures Robust Photosynthesis. Frontiers in Plant Science 9, 1617.

Smirnoff N. 2000. Ascorbate biosynthesis and function in photoprotection. Philosophical Transactions of the Royal Society B 355, 1455-1464.

Sohag AAM, Tahjib-Ul-Arif M, Polash MAS, Chowdhury MB, Afrin S, Burritt DJ, Murata Y, Hossain MA, Hossain MA. 2020. Exogenous glutathione-mediated drought stress tolerance in rice (Oryza sativa L.) is associated with lower oxidative damage and favorable ionic homeostasis. Iranian Journal of Science and Technology, Transaction A: Science 44(4), 955-971.

Sonoike K. 2011. Photo inhibition of photosystem I. Physiologia Plantarum 142, 56-64.

South PF, Cavanagh AP, Liu HW, Ort DR. 2019. Synthetic glycolate metabolism pathways stimulate crop growth and productivity in the field. Science 363, 6422.

Stephen G. Cessana, Barbara Demming A, Willium WA. 2010. Exploring Photosynthesis and Plant Stress Using Inexpensive Chlorophyll Fluorometers.

Stock D, Leslie AGW, Walker JE. 1999. Molecular architecture of the rotary motor in ATP synthase. Science 286, 1700-1705.

Suorsa M. 2015. Cyclic electron flow provides acclimatory plasticity for the photosynthetic machinery under various environmental conditions and developmental stages. Frontiers in Plant Science 6, 800.

Tahjib-Ul-Arif M, Afrin S, Polash MAS, Akter T, Ray SR, Hossain MT, Hossain MA. 2019. Role of exogenous signaling molecules in alleviating salt-induced oxidative stress in rice (Oryza sativa L.): a comparative study. Acta Physiologia Plantarum 41(5), 1-14.

TahjibUlArif M, Sohag AAM, Mostofamg, Polash MAS, Mahamud ASU, Afrin S, Hossain A, Hossain MA, Murata Y, Phan Tran LS. 2020. Comparative effects of ascobin and glutathione on copper homeostasis and oxidative stress metabolism in mitigation of copper toxicity in rice, Plant Biology.

Takagi D, Amako K, Hashiguchi M, Fukaki H, Ishizaki K, Goh T, Fukao Y, Sano R, Kurata T, Demura T, Sawa S. 2017. Chloroplastic ATP synthase builds up a proton motive force preventing production of reactive oxygen species in photosystem I. The Plant Journal 91(2), 306-324.

Takagi D, Miyake C. 2018. Proton Gradient Regulation 5supports linear electron flow to oxidize photosystem I. Physiologia Plantarum 164, 337-348.

Takagi D, Takumi S, Hashiguchi M, Sejima T, Miyake C. 2016b. Superoxide and singlet oxygen produced within the thylakoid membranes bothcause photosystem I photo inhibition. Plant Physiology 171, 1626-1634.

Taniguchi Y, Ohkawa H, Masumoto C, Fukuda T, Tamai T, Lee K, Sudoh S, Tsuchida H, Sasaki H, Fukayama H, Miyao M. 2008. Overproduction of C4photosynthetic enzymes in transgenic rice plants: anapproach to introduce the C4-like photosynthetic pathway into rice. Journal of Experimental Botany 59, 1799-1809.

Tavano CL, Donohue TJ. 2006. Development of the bacterial photosynthetic apparatus. Current Opinion in Microbiology 9(6), 625-631.

Terletskaya NV, Shcherban AB, Nesterov MA, Perfilev RN, Salina EA, Altayeva NA, Blavachinskaya IV. 2020. Drought Stress Tolerance and Photosynthetic Activity of Alloplasmic Lines T. dicoccum and T. aestivum. International Journal of Molecular Science 21(9), 3356.

Tikkanen M, Grebe S. 2018. Switching off photoprotection of photosystem I – a novel tool for gradual PSI photo inhibition. Physiologia Plantarum 162(2), 156-161.

Ting IP. 1985. Crassulacean acid metabolism. Annual Review of Plant Physiology 36(1), 595-622.

Tommos C, Babcock GT. 1999. Oxygen production in nature: A light-driven metallo radical enzyme process. Accounts of Chemical Research 37, 18-25.

Tsuchida H, Tamai T, Fukayama H, Agarie S, Nomura M, Onodera H, Ono K, Nishizawa Y, Lee BH, Hirose S, Toki S, Ku MS, Matsuoka M, Miyao M. 2001. High level expression of C4-specific NADP-malic enzyme in leaves and impairment of photoautotrophic growth in a C3 plant, rice. Plant Cell Physiology 42, 138-145.

Tyystjärvi E, Aro EM. 1996. The rate constant of photo inhibition, measured in lincomycin-treated leaves, is directly proportional to light intensity. Proceedings of the National Academy of Sciences 93(5), 2213-2218.

Tyystjärvi E. 2008. Photoinhibition of photosystem II and photo damage of the oxygen evolving manganese cluster. Coordination Chemistry Review 252(3-4), 361-376.

Vass I, Styring S, Hundal T, Koivuniemi A, Aro E, Andersson B. 1992. Reversible and irreversible intermediates during photoinhibition of photosystem II: stable reduced QA species promote chlorophyll triplet formation. Proceedings of the National Academy of Sciences 89(4), 1408-1412.

Von Caemmerer S, Furbank RT. 2003. The C4 pathway: an efficient CO2 pump. Photosynthesis Research 77, 191-207.

Von Caemmerer S, Ghannoum O, Furbank RT. 2017. C4 photosynthesis: 50 years of discovery and innovation. Journal of Experimental Botany 68(2), 97-102.

Von Caemmerer S, Quick WP, Furbank RT. 2012. The development of C4 rice: current progress and future challenges. Science 336, 1671-1672.

Wang P, Fouracre J, Kelly S, Karki S, Gowik U, Aubry S, Shaw MK, Westhoff P, Slamet-Loedin IH, Quick WP, Hibberd JM. 2013. Evolution of GOLDEN2-LIKE gene function in C3 and C4 plants. Planta 237(2), 481-495.

Wang P, Vlad DL, Langdale JA. 2016. Finding the genes to build Crice. Current Opinion in Plant Biology 31, 44-50.

Wang S, Tholen D, Zhu XG. 2017. C4 photosynthesis in C3 rice: a theoretical analysis of biochemical and anatomical factors. Plant Cell Environment 40(1), 80-94.

Weber APM, von Caemmerer S. 2010. Plastid transport and metabolism of C3 and C4 plants: comparative analysis and possible biotechnological mexploitation. Current Opinion in Plant Biology 13, 257-265.

Whatley FT, Allen MB. 1954. Photosynthesis by isolated chloroplasts. II. Photosynthetic phosphorylation, the conversion of light into phosphate-bond energy. The Journal of the American Chemical Society 76, 6324-6328.

Whitmarsh J. 1999. The photosynthetic process, In Concepts in photobiology. Springer, Dordrecht 11-51.

Wingler A, Lea PJ, Leegood RC. 1999. Photorespiratory metabolism of glyoxylate and formate in glycine-accumulating mutants of barley and Amaranthus edulis. Planta 207(4), 518-526.

Yamamoto H Y, Nakayama T, Chichester CO. 1962. Studies on the light and dark inter conversions of leaf xanthophylls. Archives of Biochemistry and Biophysics 97(1), 168-173.

Yokthongwattana K, Melis A. 2008. Photo inhibition and Recovery in Oxygenic Photosynthesis: Mechanism of a Photosystem II Damageand Repair Cycle. In: Demmig-Adams, B., Adams, W. W. III., and Mattoo, A. K., (Eds), Photo protection, Photo inhibition, Gene Regulation, and Environment. Springer, Dordrecht 175-191.

Zamani-Nour S, Lin HC, Walker BJ, Mettler-Altmann T, Khoshravesh R, Karki S, Bagunu E, Sage TL, Quick WP, Weber APM. 2020. Overexpression of the chloroplastic 2-oxoglutarate/malate transporter disturbs carbon and nitrogen homeostasis in rice. Journal of Experimental Botany 72(1), 137-152.

Zheng XT, Chen YL, Zhang XH, CaimL, Yu ZC, Peng CL. 2019. ANS-deficient Arabidopsis is sensitive to high light due to impaired anthocyanin photo protection. Functional Plant Biology 46(8), 756-765.

Ziehe D, Dünschede B, Schünemann D. 2018. Molecular mechanism of SRP-dependent light-harvesting protein transport to the thylakoid membrane in plants. Photosynthesis Research 138(3), 303-313.

Zivcak M, Brestic M, Kunderlikova K, Sytar O, Allakhverdiev SI. 2015b. Repetitive light pulse-induced photoinhibition of photosystem I severely affects CO2 assimilation and photoprotection in wheat leaves. Photosynthesis Research 126, 449-463.