植物光系统的结构与功能
Structure and Function of Plant Light System
DOI: 10.12677/BR.2019.82023, PDF,  被引量   
作者: 葛倩雯, 金宝花：浙江师范大学，化学与生命科学学院，浙江 金华
关键词: 光合作用；光系统I；光系统II；天线系统；Photosynthesis； Photosystem I； Photosystem II； Antenna System
摘要: 通过将光能转化为化学能并产生分子氧和消耗二氧化碳，光合作用氧合成对于地球生命的发展和维持是必不可少的。后一个过程负责将二氧化碳从原始大气中的极高水平降低到目前的低水平，从而将全球温度降低到有利于生命发展的水平。光系统I和光系统II是两种多蛋白复合物，其含有收获光子所必需的颜料，并利用光能催化产生高能化合物的主要光合作用的反应。两种光系统都是由核心复合物组成的高度有组织的膜超复合物，包含启动电子传递的反应中心，以及对于光捕获和光合作用活动调节很重要的外围天线系统。一方面两个光系统催化的化学反应和它们的详细结构都不同，另一方面它们有许多相似之处。在本综述中，我们对光系统I和光系统II的结构和功能进行详细的归纳和描述。
Abstract: Photosynthetic oxygen synthesis is essential for the development and maintenance of life on Earth by converting light energy into chemical energy and producing molecular oxygen and carbon dioxide. The latter process is responsible for reducing carbon dioxide from extremely high levels in the original atmosphere to current low levels, thereby reducing global temperatures to levels conducive to life. Photosystem I and Photosystem II are two multiprotein complexes that contain the pigments necessary to harvest photons and utilize photo energetic catalysis to produce the main photosynthesis reactions of high energy compounds. Both photosystems are highly organized membrane super complexes composed of core complexes, including reaction centers that initiate electron transport, and peripheral antenna systems that are important for light harvesting and photosynthesis activity regulation. On the one hand, the two photosystems catalyze chemical reactions and their detailed structures are different; on the other hand they have many similarities. In this review, we present and describe in detail the structure and function of Photosystem I and Photosystem II.
文章引用：葛倩雯, 金宝花. 植物光系统的结构与功能[J]. 植物学研究, 2019, 8(2): 170-179. https://doi.org/10.12677/BR.2019.82023

参考文献

[1] Bekker, A., Holland, H.D., Wang, P.L., Rumble, D., Stein, H.J., Hannah, J.L., Coetzee, L.L. and Beukes, N.J. (2004) Dating the Rise of Atmospheric Oxygen. Nature, 427, 117-120. [Google Scholar] [CrossRef] [PubMed]
[2] Hohmann-Marriott, M.F. and Blankenship, R.E. (2011) Evolution of Photosynthesis. Annual Review of Plant Biology, 62, 515-548. [Google Scholar] [CrossRef] [PubMed]
[3] Hasan, S.S., Yamashita, E., Ryan, C.M., White-legge, J.P. and Cramer, W.A. (2011) Conservation of Lipid Functions in Cytochrome BC Complexes. Journal of Molecular Biology, 414, 145-162. [Google Scholar] [CrossRef] [PubMed]
[4] Paul, M. (2013) Photosynthesis. Plastid Biology, Energy Conversion and Carbon Assimilation. Annals of Botany, 99, 972.
[5] Horton, P., Ruban, A.V. and Walters, R.G. (1996) Regulation of Light Harvesting in Green Plants. Annual Review of Plant Physiology and Plant Molecular Biology, 47, 655-684. [Google Scholar] [CrossRef] [PubMed]
[6] Jansson, S. (2006) Photoprotection, Photoinhibition, Gene Regulation, and Environment. In: Demmig-Adams, B., Adams III, W.W. and Mattoo, A.K., Eds., Advances in Photosynthesis and Respiration, Vol. 21, Springer, Dordrecht, 145-153.
[7] Golbeck, J.H. (1994) The Molecular Biology of Cyanobacteria. In: Bryant, D.A., Ed., Advances in Photosynthesis: Vol. 1, Kluwer Academic Publishers, Dordrecht, 319-360.
[8] Chitnis, P.R. (1996) Photosystem I. Plant Physiology, 111, 661-669. [Google Scholar] [CrossRef] [PubMed]
[9] Döring, G., Bailey, J.L., Kreutz, W., et al. (1968) Some New Results in Photosynthesis. Die Naturwissenschaften, 55, 219-224. [Google Scholar] [CrossRef] 
[10] Norris, J.R., Uphaus, R.A., Crespi, H.L., et al. (1971) Electron Spin Resonance of Chlorophyll and the Origin of Signal I in Photosynthesis. Proceedings of the National Academy of Sciences, 68, 625-628. [Google Scholar] [CrossRef] [PubMed]
[11] Jordan, P., Fromme, P., Witt, H.T., et al. (2001) Three-Dimensional Structure of Cyanobacterial Photosystem I at 2.5 Å Resolution. Nature, 411, 909. [Google Scholar] [CrossRef] [PubMed]
[12] Ben-Shem, A., Frolow, F. and Nelson, N. (2003) Crystal Structure of Plant Photosystem I. Nature, 426, 630-635. [Google Scholar] [CrossRef] [PubMed]
[13] Schubert, W.D., Klukas, O., Saenger, W., et al. (1998) A Common Ancestor for Oxygenic and Anoxygenic Photosynthetic Systems: A Comparison Based on the Structural Model of Photosystem I. Journal of Molecular Biology, 280, 297-314. [Google Scholar] [CrossRef] [PubMed]
[14] Fromme, P., Schubert, W.D. and Krauß, N. (1994) Structure of Photosystem I: Suggestions on the Docking Sites for Plastocyanin, Ferredoxin and the Coordination of P700. Biochimica et Biophysica Acta (BBA)-Bioenergetics, 1187, 99-105. ttps://doi.org/10.1016/0005-2728(94)90092-2
[15] Deisenhofer, J., Epp, O., Sinning, I., et al. (1995) Crystallofraphic Refinement at 2.3-Angstrom Resolution and Refined Model of the Photosynthetic Reaction Centre from Phodopseudomonas Viridis. Journal of Molecular Biology, 246, 429-457. [Google Scholar] [CrossRef] [PubMed]
[16] Rhee, K.H. and Morris, E.P. (1998) Three Dimensional Structure of the Photosystem II Reaction Centre at 8 Å. Nature, 396, 283-286. [Google Scholar] [CrossRef] [PubMed]
[17] Zouni, A., Witt, H.T., Kern, J., et al. (2001) Crystal Structure of Photosystem II from Synechococcus elongatus at 3.8 A Resolution. Nature, 409, 739-743. [Google Scholar] [CrossRef] [PubMed]
[18] Hippler, M., Reichert, J., Sutter, M., et al. (1996) The Plastocyanin Binding Domain of Photosystem I. EMBO Journal, 15, 6374-6384. [Google Scholar] [CrossRef] [PubMed]
[19] Hoj, P.B., Svendsen, I., Scheller, H.V., et al. (1987) Identification of a Chloroplast-Encoded 9-kDa Polypeptide as a 2[4Fe-4S] Protein Carrying Centers A and B of Photosystem I. Journal of Biological Chemistry, 262, 12676-12684.
[20] Golbeck, J.H. (1992) Structure and Function of Photosystem I. Annual Review of Plant Physiology and Plant Molecular Biology, 43, 3204-3210. [Google Scholar] [CrossRef] 
[21] Lelong, C., Boekema, E.J., Kruip, J., et al. (1996) Characterization of a Redox Active Cross-Linked Complex between Cyanobacterial Photosystem I and Soluble Ferredoxin. EMBO Journal, 15, 2160-2168. [Google Scholar] [CrossRef] [PubMed]
[22] Chitnis, V.P., Jung, Y.S., Albee, L., et al. (1996) Mutational Analysis of Photosystem I Polypeptides Role of PsaD and the Lysyl 106 Redesidue in the Reductase Activity of Photosystem I. Journal of Biological Chemistry, 271, 11772-11780. [Google Scholar] [CrossRef] [PubMed]
[23] Chitnis, V.P., Ke, A. and Chitnis, P.R. (1997) The PsaD Subunit of Photosystem I. Mutations in the Basic Domain Reduce the Level of PsaD in the Membranes. Plant Physiology, 115, 1699-1705. [Google Scholar] [CrossRef] [PubMed]
[24] Barth, P., Lagoutte, B. and Sétif, P. (1998) Ferredoxin Reduction by Photosystem I from Synechocystis sp. PCC 6803: Toward an Understanding of the Respective Roles of Subunits PsaD and PsaE in Ferredoxin Binding. Biochemistry, 37, 16233-16241. [Google Scholar] [CrossRef] [PubMed]
[25] Pandini, V., Alessandro Aliverti, A. and Zanetti, G. (1999) Interaction of the Soluble Recombinant PsaD Subunit of Spinach Photosystem I with Ferredoxin I. Biochemistry, 38, 10707-10713. [Google Scholar] [CrossRef] [PubMed]
[26] Sonoike, K., Hatanaka, H. and Katoh, S. (1993) Small Subunits of Photosystem I Reaction Center Complexes from Synechococcus elongatus. II. The PsaE Gene Product Has a Role to Promote Interaction between the Terminal Electron Acceptor and Ferredoxin. Biochimica et Biophysica Acta (BBA)—Bioenergetics, 1141, 52-57. [Google Scholar] [CrossRef] 
[27] Weber, N. and Strotmann, H. (1993) On the Function of Subunit PsaE in Chloroplast Photosystem I. Biochimica et Biophysica Acta, 1143, 204-210. [Google Scholar] [CrossRef] [PubMed]
[28] Rousseau, F., Sétif, P. and Lagoutte, B. (1993) Evidence for the Involvement of PSI-E Subunit in the Reduction of Ferredoxin by Photosystem I. EMBO Journal, 12, 1755-1765. [Google Scholar] [CrossRef] [PubMed]
[29] Zhao, J., Snyder, W.B., Mühlenhoff, U., et al. (2010) Cloning and Characterization of the PsaE Gene of the Cyanobacterium Synechococcus sp. PCC 7002: Characterization of a PsaE Mutant and Overproduction of the Protein in Escherichia coli. Molecular Microbiology, 9, 183-194. [Google Scholar] [CrossRef] [PubMed]
[30] Mühlenhoff, U., Haehnel, W., Witt, H., et al. (1993) Genes Encoding Eleven Subunits of Photosystem I from the Thermophilic Cyanobacterium Synechococcus sp. Gene, 127, 71-78. [Google Scholar] [CrossRef] 
[31] Schluchter, W.M., Shen, G., Zhao, J., et al. (2010) Characterization of PsaI and PsaL Mutants of Synechococcus sp. Strain PCC 7002: A New Model for State Transitions in Cyanobacteria. Photochemistry & Photobiology, 64, 53-66. [Google Scholar] [CrossRef] [PubMed]
[32] Xu, Q., Dan, H., Chitnis, V.P., et al. (1995) Mutational Analysis of Photosystem I Polypeptides in the Cyanobacterium Synechocystis sp. PCC 6803. Targeted Inactivation of PsaI Reveals the Function of PsaI in the Structural Organization of PsaL. Journal of Biological Chemistry, 270, 16243-16250. [Google Scholar] [CrossRef] [PubMed]
[33] Fromme, P. (1996) Structure and Function of Photosystem I. Current Opinion in Structural Biology, 6, 473-484. [Google Scholar] [CrossRef] 
[34] Armbrust, T.S., Chitnis, P.R. and Guikema, J.A. (1996) Organization of Photosystem I Polypeptides Examined by Chemical Cross-Linking. Plant Physiology, 111, 1307-1312. [Google Scholar] [CrossRef] [PubMed]
[35] Fischer, N., Boudreau, E., Hippler, M., et al. (1999) A Large Fraction of PsaF Is Nonfunctional in Photosystem I Complexes Lacking the PsaJ Subunit. Biochemistry, 38, 5546-5552. [Google Scholar] [CrossRef] [PubMed]
[36] Anandan, S., Vainstein, A. and Thornber, J.P. (1989) Correlation of Some Published Amino Acid Sequences for Photosystem I Polypeptides to a 17 kDa LHCI Pigment-Protein and to Subunits III and IV of the Core Complex. Febs Letters, 256, 150-154. [Google Scholar] [CrossRef] [PubMed]
[37] Kjaerulff, S., Andersen, B., Nielsen, V.S., et al. (1993) The PSI-K Subunit of Photosystem I from Barley (Hordeum vulgare L.). Evidence for a Gene Duplication of an Ancestral PSI-G/K Gene. Journal of Biological Chemistry, 268, 18912-18916.
[38] Jansson, S., Andersen, B. and Scheller, H.V. (1996) Nearest-Neighbor Analysis of Higher-Plant Photosystem I Holocomplex. Plant Physiology, 112, 409-420. [Google Scholar] [CrossRef] [PubMed]
[39] Klimmek, F., Ganeteg, U., Jansson, S., et al. (2005) Structure of the Higher Plant Light Harvesting Complex I:  In Vivo Characterization and Structural Interdependence of the Lhca Proteins. Biochemistry, 44, 3065-3073. [Google Scholar] [CrossRef] [PubMed]
[40] Croce, R., Mozzo, M., Morosinotto, T., et al. (2007) Singlet and Triplet State Transitions of Carotenoids in the Antenna Complexes of Higher-Plant Photosystem I. Biochemistry, 46, 3846-3855. [Google Scholar] [CrossRef] [PubMed]
[41] Croce, R., Morosinotto, T., Castelletti, S., et al. (2002) The Lhca Antenna Complexes of Higher Plants Photosystem I. Biochimica et Biophysica Acta, 1556, 29-40. [Google Scholar] [CrossRef] 
[42] Amunts, A., Drory, O. and Nelson, N. (2007) The Structure of a Plant Photosystem I Supercomplex at 3.4 Å Resolution. Nature, 447, 58-63. [Google Scholar] [CrossRef] [PubMed]
[43] Morosinotto, T., Breton, J., Bassi, R., et al. (2003) The Nature of a Chlorophyll Ligand in Lhca Proteins Determines the Far Red Fluorescence Emission Typical of Photosystem I. Journal of Biological Chemistry, 278, 49223-49229. [Google Scholar] [CrossRef] 
[44] Croce, R., Chojnicka, A., Morosinotto, T., et al. (2007) The Low-Energy Forms of Photosystem I Light-Harvesting Complexes: Spectroscopic Properties and Pigment-Pigment Interaction Characteristics. Biophysical Journal, 93, 2418-2428. [Google Scholar] [CrossRef] [PubMed]
[45] Schmid, V.H.R., et al. (2002) Pigment Binding of Photosystem I Light-Harvesting Proteins. Journal of Biological Chemistry, 277, 37307-37314. [Google Scholar] [CrossRef] 
[46] Croce, R., Zucchelli, G., Garlaschi, F.M., et al. (1996) Excited State Equilibration in the Photosystem I? Light-Harvesting I Complex: P700 Is Almost Isoenergetic with Its Antenna. Biochemistry, 35, 8572-8579. [Google Scholar] [CrossRef] [PubMed]
[47] Wientjes, E. and Croce, R. (2011) The Light-Harvesting Complexes of Higher-Plant Photosystem I: Lhca1/4 and Lhca2/3 form Two Red-Emitting Heterodimers. Biochemical Journal, 433, 477-485. [Google Scholar] [CrossRef] 
[48] Albertsson, P. (2001) A Quantitative Model of the Domain Structure of the Photosynthetic Membrane. Trends in Plant Science, 6, 349-354. [Google Scholar] [CrossRef] 
[49] Dekker, J.P. and Boekema, E.J. (2005) Supramolecular Organization of Thylakoid Membrane Proteins in Green Plants. BBA Bioenergetics, 1706, 12-39.
[50] Shi, L.X., Hall, M., Funk, C., et al. (2012) Photosystem II, a Growing Complex: Updates on Newly Discovered Components and Low Molecular Mass Proteins. Biochimica et Biophysica Acta, 1817, 13-25. [Google Scholar] [CrossRef] [PubMed]
[51] Shi, L.X. and Schröder, W.P. (2004) The Low Molecular Mass Subunits of the Photosynthetic Supracomplex, Photosystem II. Biochimica et Biophysica Acta, 1608, 75-96.
[52] Zouni, A., Kern, J., Loll, B., et al. (2001) Biochemical Characterization and Crystal Structure of Water Oxidizing Photosystem II from Synechococcus elongatus. Science Access, 3.
[53] Kamiya, N. and Shen, J.R. (2003) Crystal Structure of Oxygen-Evolving Photosystem II from Thermo Synechococcus vulcanus at 3.7 Å Resolution. Proceedings of the National Academy of Sciences of the United States of America, 100, 98-103. [Google Scholar] [CrossRef] [PubMed]
[54] Stefano, C., Tania, T., Robert, J., et al. (2014) A Comparison between Plant Photosystem I and Photosystem II Architecture and Functioning. Current Protein & Peptide Science, 15, 296-331. [Google Scholar] [CrossRef] [PubMed]
[55] Peter, G.F. and Thornber, J.P. (1991) Biochemical Composition and Organization of Higher Plant Photosystem II Light-Harvesting Pigment-Proteins. Journal of Biological Chemistry, 266, 16745-16754.
[56] Standfuss, J.A., Lamborghini, M. and Kühlbrandt, W. (2005) Mechanisms of Photoprotection and Nonphotochemical Quenching in Pea Light Harvesting Complex at 2.5 Å Resolution. EMBO Journal, 24, 919-928. [Google Scholar] [CrossRef] [PubMed]
[57] Kuhlbrandt, W., Wang, D.N. and Fujiyoshi, Y. (1994) Atomic Model of Plant Light-Harvesting Complex by Electron Crystallography. Nature, 367, 614-621. [Google Scholar] [CrossRef] [PubMed]
[58] Wei, X., Su, X., Cao, P., et al. (2016) Structure of Spinach Photosystem II-LHCII Supercomplex at 3.2Å Resolution. Nature, 534, 69. [Google Scholar] [CrossRef] [PubMed]
[59] Su, X., Ma, J., Wei, X., et al. (2017) Structure and Assembly Mechanism of Plant C2S2M2-Type PSII-LHCII Supercomplex. Science, 357, 815-820. [Google Scholar] [CrossRef] [PubMed]
[60] Van Bezouwen, L.S., Caffarri, S., Kale, R.S., et al. (2017) Subunit and Chlorophyll Organization of the Plant Photosystem II Supercomplex. Nature Plants, 3, Article No. 17080. [Google Scholar] [CrossRef] [PubMed]
[61] Labate, M.T., Ko, K., Ko, Z.W., et al. (2004) Constitutive Expression of Pea Lhcb 1-2 in Tobacco Affects Plant Development, Morphology and Photosynthetic Capacity. Plant Molecular Biology, 55, 701-714. [Google Scholar] [CrossRef] [PubMed]
[62] Jenny, A., Mark, W., Walters, R.G., et al. (2003) Absence of the Lhcb1 and Lhcb2 Proteins of the Light-Harvesting Complex of Photosystem II—Effects on Photosynthesis, Grana Stacking and Fitness. The Plant Journal, 35, 350-361. [Google Scholar] [CrossRef] 
[63] Garab, G. (2014) Hierarchical Organization and Structural Flexibility of Thylakoid Membranes. BBA—Bioenergetics, 1837, 481-494. [Google Scholar] [CrossRef] [PubMed]
[64] Garab, G. (2016) Self-Assembly and Structural-Functional Flexibility of Oxygenic Photosynthetic Machineries: Personal Perspectives. Photosynthesis Research, 127, 131-150. [Google Scholar] [CrossRef] [PubMed]
[65] Liu, Z., Yan, H., Wang, K., et al. (2004) Crystal Structure of Spinach Major Light-Harvesting Complex at 2.72 Å Resolution. Nature, 428, 287-292. [Google Scholar] [CrossRef] [PubMed]
[66] Pan, X., Li, M., Wan, T., et al. (2011) Structural Insights into Energy Regulation of Light-Harvesting Complex CP29 from Spinach. Nature Structural & Molecular Biology, 18, 309-315. [Google Scholar] [CrossRef] [PubMed]