叶绿素含量降低对水稻叶片光抑制与光合电子传递的影响

doi:10.3864/j.issn.0578-1752.2016.19.004

摘要/Abstract

摘要： 【目的】水稻突变体叶片叶绿素含量显著低于其野生型，但其光合电子传递效率与净光合值显著高于对照，文章旨在阐明其生理学机理并探讨其在高光效育种中的潜在应用价值。【方法】通过人工气候室的盆栽试验设置高光强（光照强度为700—800 µmol·m^-2·s^-1）与低光强（光照强度约为100—200 µmol·m^-2·s^-1）2个处理，并结合大田试验，观测突变体与野生型的叶绿素荧光、超氧阴离子含量、丙二醛含量、SOD酶活性、叶绿体超微结构、叶片荧光显微结构与冠层温度。【结果】突变体材料叶绿素含量显著低于其野生型，并且光照强度对其叶绿素含量的效应也不相同。随着光照强度的增强，突变体叶片叶绿素含量增加60%，而野生型品种叶片叶绿素含量却降低20%以上。光反应曲线表明，突变体光合值显著高于野生型，尤其在1 000 µmol·m^-2·s^-1光照下，低光强与高光强处理中低叶绿素含量突变体光合值分别比野生型高9.4%和46.5%。叶绿素荧光数据也表明，水稻突变体的电子传递速率（ETR）、光系统Ⅱ的量子产量（Φ_PSⅡ）、光化学淬灭（qP）、光下最大光化学效率（F′v/F′m）显著高于其野生型。在高光强处理下，野生型材料中的应激活性氧超氧阴离子与丙二醛（MDA）含量受到光抑制。叶绿体超微结构与荧光显微结构表明，野生型材料叶绿体受到一定程度的损伤，且其维管束间距离显著大于突变体，其维管束面积小于突变体，不利于水分在叶片中的传输。田间冠层热力学图像表明中午高温、高光照条件下，突变体冠层温度显著低于其野生型。综合以上结果，野生型叶片叶绿素含量高，在高光强下会导致过量光吸收，光系统Ⅱ电子传递效率下降，超氧阴离子与丙二醛累计，导致叶绿素结构与功能的破坏，因此，其光合值显著低于突变体材料。同时过量光吸收会导致叶片与冠层温度上升，不利于冠层群体光合。【结论】在未来高光效育种中选育叶绿素含量适当降低的品种，有助于避免高光强下叶片的过量光吸收，从而缓解活性氧的产生与光抑制，并有利于降低冠层温度从而缓解水稻群体光合“午休”现象。

关键词: 水稻, 光合速率, 叶绿素, 叶绿素荧光, 光抑制

Abstract: 【Objective】The chlorophyll content of a chlorophyll-deficit rice mutant (YL) is significantly lower than its wild type (WT), but its photosynthetic electron transport rate and net photosynthetic rate are significantly higher than its WT. The aim of this study is to understand the physiological basis, and its potential use in high photosynthetic efficiency breeding was prospected.【Method】A pot experiment in the climate chamber at high light intensity (HL) and low light intensities (LL), and a field experiment were conducted. Chlorophyll fluorescence, the concentration of super oxygen anion and malodialdehyde, superoxide dismutase activity, light fluorescent and electron micrographs, canopy temperature were investigated. 【Result】 The results show that the chlorophyll content was significantly lower in the mutant than its WT, and the light treatment had different effects on chlorophyll content in the mutant and its WT. When compared the HL and LL treatments, the chlorophyll content was increased by 60% in the mutant, but decreased by 20% in the WT. The light response curves showed that the mutant has a higher photosynthetic capacity than its WT. At the irradiance of 1 000 µmol·m^-2·s^-1, the photosynthetic rate was 46.5% and 9.4% higher in the mutant than its WT, in HL and LL treatments, respectively. The chlorophyll fluorescence measurements indicate that the photosynthetic electron transport rate (ETR), photosystem Ⅱ efficiency (Φ_PS_Ⅱ), photosynthetic quenching (qP), and maximum efficiency of open photosystem Ⅱ in the light (F′v/F′m) were significantly higher in mutant than its WT. The content of oxygen anion and malondialdehyde were higher in WT than the mutant, especially in HL treatment, indicating photoinhibition in WT. The electron micrographs and light fluorescent micrographs of the mutant and its WT indicated that the chloroplast was to some extent harmed. The inter-vein distance was also found larger in WT than in the mutant, and the area of vascular bundle is smaller in WT than in the mutant, indicating a better water status in mutant than in its WT. The thermal image indicated that the canopy temperature was significantly lower in mutant than in its WT at noon with high irradiance. All the results suggested that the high chlorophyll content in the WT excessively absorbed light energy, causing photoinhibition (high oxygen anion and malondialdehyde content, reduced photosystem Ⅱefficiency and decreased SOD activity), and had a lower photosynthetic rate than the mutant. The excessively absorbed light energy also contributed the higher canopy temperature in WT than in mutant, which is adverse for canopy photosynthesis.【Conclusion】All these results implicate that selecting for moderate chlorophyll content in breeding would help avoiding the generation of reactive oxygen species and alleviating the photoinhibition, improving photosynthesis, especially at noon under high solar radiation.

Key words: rice, photosynthetic rate, chlorophyll, chlorophyll fluorescence, photoinhibition

周振翔，李志康，陈颖，王志琴，杨建昌，顾骏飞. 叶绿素含量降低对水稻叶片光抑制与光合电子传递的影响[J]. 中国农业科学, 2016, 49(19): 3709-3720.

ZHOU Zhen-xiang, LI Zhi-kang, CHEN Ying, WANG Zhi-qin, YANG Jian-chang, GU Jun-fei. Effects of Reduced Chlorophyll Content on Photoinhibition and Photosynthetic Electron Transport in Rice Leaves[J]. Scientia Agricultura Sinica, 2016, 49(19): 3709-3720.

0
/ / 推荐

导出引用管理器 EndNote|Reference Manager|ProCite|BibTeX|RefWorks

链接本文: https://www.chinaagrisci.com/CN/10.3864/j.issn.0578-1752.2016.19.004

https://www.chinaagrisci.com/CN/Y2016/V49/I19/3709

参考文献

[1] Murchie E H, Pinto M, Horton P. Agriculture and the new challenges for photosynthesis research. New Phytologist, 2009, 181(3): 532-552.

[2] Peng S, Khush G S, Virk P, Tang Q, Zou Y. Progress in ideotype breeding to increase rice yield potential. Field Crops Research, 2008, 108: 32-38.

[3] Long S P. We need winners in the race to increase photosynthesis in rice, whether from conventional breeding, biotechnology or both. Plant, Cell & Environment, 2014, 37(1): 19-21.

[4] Long S P, Marshall-Colon A, Zhu X G. Meeting the global food demand of the future by engineering crop photosynthesis and yield potential. Cell, 2015, 161(1): 56-66.

[5] von Caemmerer S, Quick W P, Furbank R T. The development of C4 rice: current progress and future challenges. Science, 2012, 336(6089): 1671-1672.

[6] Krause G H, Weis E. Chlorophyll fluorescence and photosynthesis: the basics. Annual Review of Plant Biology, 1991, 42(1): 313-349.

[7] Murchie E H, Niyogi K K. Manipulation of photoprotection to improve plant photosynthesis. Plant Physiology, 2011, 155: 86-92.

[8] Zhu X G, Long S P, Ort D R. Improving photosynthetic efficiency for greater yield. Annual Review of Plant Biology, 2010, 61: 235-261.

[9] Demao J, Xia L, Xueqing H, WEI C, TINGYUN K, MAURICE S B K. The characteristics of CO₂ assimilation of photosynthesis and chlorophyll fluorescence in transgenic PEPC rice. Chinese Science Bulletin, 2001, 46(13): 1080-1084.

[10] Ort D R, Merchant S S, Alric J, Barkan A, Blankenship R E, Bock R, Croce R, Hanson M R, Hibberd J M, Long S P. Redesigning photosynthesis to sustainably meet global food and bioenergy demand. Proceedings of the National Academy of Sciences of the USA, 2015, 112: 8529-8536.

[11] Ort D R, Zhu X G, Melis A. Optimizing antenna size to maximize photosynthetic efficiency. Plant Physiology, 2011, 155: 79-85.

[12] Nakajima Y, Itayama T. Analysis of photosynthetic productivity of microalgal mass cultures. Journal of Applied Phycology, 2003, 15: 497-505.

[13] Kirst H, Formighieri C, Melis A. Maximizing photosynthetic efficiency and culture productivity in cyanobacteria upon minimizing the phycobilisome light-harvesting antenna size. Biochimica et Biophysica Acta, 2014, 1837:1653-1664.

[14] Holden M. Chlorophyll// Goodwin T W. Chemistry and Biochemistry of Plant Pigments.2nd ed. London: Academic Press, 1976: 1-37.

[15] Genty B, Briantais J M, Baker N R. The relationship between the quantum yield of photosynthetic electron transport and quenching of chlorophyll fluorescence. Biochimica et Biophysica Acta, 1989, 990: 87-92.

[16] Maxwell K, Johnson G N. Chlorophyll fluorescence-A practical guide. Journal of Experimental Botany, 2000, 51: 659-668.

[17] RODRÍGUEZ-SERRANO M, ROMERO-PUERTAS M C, Zabalza A N A, CORPAS F J, GÓMEZ M, DEL RÍO L A, SANDALIO L M. Cadmium effect on oxidative metabolism of pea (Pisum sativum L.) roots. Imaging of reactive oxygen species and nitric oxide accumulation in vivo. Plant, Cell & Environment, 2006, 29(8): 1532-1544.

[18] 王爱国, 罗广华, 邵从本, 吴淑君, 郭俊彦. 大豆种子超氧物歧化酶的研究. 植物生理学报, 1983, 9(1): 77-84.

Wang A G, Luo G H, Shao C B, Wu S J, Guo J Y. A study on the superoxide dismutase of soybean seeds. Acta Phytotaxonomica Sinica, 1983, 9(1): 77-84. (in Chinese)

[19] 赵世杰, 许长成, 邹琦, 孟庆伟. 植物组织中丙二醛测定方法的改进. 植物生理学通讯, 1994, 30: 207-210.

Zhao S J, Xu C C, Zou Q, Meng Q W. Improvements of method for measurement of malondialdehyde in plant tissues. Plant Physiology Communications, 1994, 30: 207-210. (in Chinese)

[20] Zhao H, Kalivendi S, Zhang H, JOSEPH J, NITHIPATIKOM K, Vásquez-Vivar J, Kalyanaraman B. Superoxide reacts with hydroethidine but forms a fluorescent product that is distinctly different from ethidium: potential implications in intracellular fluorescence detection of superoxide. Free Radical Biology and Medicine, 2003, 34(11): 1359-1368.

[21] 刘占才, 牛俊英. 超氧阴离子自由基对生物体的作用机理研究. 焦作教育学院学报, 2002, 18(4): 48-51.

Liu Z C, Niu J Y. Research on the mechanism of the organism on superoxide anion. Journal of Jiaozuuo Institute of Education, 2002, 18(4): 48-51. (in Chinese)

[22] Horton P. Prospects for crop improvement through the genetic manipulation of photosynthesis: morphological and biochemical aspects of light capture. Journal of Experimental Botany, 2000, 51: 475-485.

[23] Li Y, Ren B, Gao L, DING L, JIANG D, XU X, SHEN Q, GUO S. Less chlorophyll does not necessarily restrain light capture ability and photosynthesis in a chlorophyll-deficient rice mutant. Journal of Agronomy and Crop Science, 2013, 199(1): 49-56.

[24] Yin X, Struik P C. Constraints to the potential efficiency of converting solar radiation into phytoenergy in annual crops: from leaf biochemistry to canopy physiology and crop ecology. Journal of Experimental Botany, 2015, 62: 3489-3499.

[25] Mu H, Jiang D, Wollenweber B, DAI T, JING Q, CAO W. Long-term low radiation decreases leaf photosynthesis, photochemical efficiency and grain yield in winter wheat. Journal of Agronomy and Crop Science, 2010, 196(1): 38-47.

[26] Geacintov N E, Breton J, Knox R S. Energy transfer and fluorescence mechanisms in photosynthetic membranes. Critical reviews in plant sciences, 1987, 5(1): 1-44.

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

[28] Foyer C H, Noctor G. Leaves in the dark see the light. Science, 1999, 284(5414): 599-601.

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

[30] Krause G H. Photoinhibition of photosynthesis. An evaluation of damaging and protective mechanisms. Physiologia Plantarum, 1988, 74(3): 566-574.

[31] Barber J. Molecular basis of the vulnerability of photosystem II to damage by light. Functional Plant Biology, 1995, 22(2): 201-208.

[32] Takagi D, Takumi S, Hashiguchi M, SEJIMA T, MIYAKE C. Superoxide and singlet oxygen produced within the thylakoid membranes both cause photosystem I photoinhibition. Plant Physiology, 2016, Doi: 10.1104/pp.16.00246.

[33] HUANG W, YANG Y J, ZHANG J L, HU H, ZHANG S B. PSI photoinhibition is more related to electron transfer from PSII to PSI rather than PSI redox state in Psychotria rubra. Photosynthesis Research, 2016, 129(1): 85-92.

[34] TIWARI A, MAMEDOV F, GRIECO M, SUORSA M, JAJOO A, STYRING S, TIKKANEN M, ARO E M. Photodamage of iron- sulphur clusters in photosystem I induces non-photochemical energy dissipation. Nature Plants, 2016, 2: 16035.

[35] Clarke J E, Johnson G N.In vivo temperature dependence of cyclic and pseudocyclic electron transport in barley. Planta, 2001, 212(5/6): 808-816.

[36] Ort D R, Merchant S S, Alric J, BARKAN A, BLANKENSHIP R E, BOCK R, CROCE R, HANSON M R, HIBBERD J M, LONG S P, MOORE T A, MORONEY J, NIYOGI K K, PARRY M A J, PERALTA-YAHYA P P, PRINCE R C, REDDING K E, SPALDING M H, VAN WIJK K J, VERMAASW F J, CAEMMERER S V, WEBWE A P M, YEATES T O, YUAN J S, ZHU X G. Redesigning photosynthesis to sustainably meet global food and bioenergy demand. Proceedings of the National Academy of Science of the USA, 2015, 112: 8529-8536.

[37] Ort D R, Zhu X G, Melis A. Optimizing antenna size to maximize photosynthetic efficiency. Plant Physiology, 2011, 155: 79-85.

[38] Polle J E, Kanakagiri S D, Melis A. tla1, a DNA insertional transformant of the green alga Chlamydomonas reinhardtii with a truncated light-harvesting chlorophyll antenna size. Planta, 2003, 217: 49-59.

[39] Melis A. Solar energy conversion efficiencies in photosynthesis: minimizing the chlorophyll antennae to maximize efficiency. Plant Science, 2009, 177: 272-280.

[40] Sukenik A, Bennett J, Falkowski P. Light-saturated photosynthesis—limitation by electron transport or carbon fixation? Biochimica et Biophysica Acta (BBA)-Bioenergetics, 1987, 891(3): 205-215.

[41] Long S P, ZHU X G, Naidu S L, ORT D R. Can improvement in photosynthesis increase crop yields? Plant, Cell & Environment, 2006, 29(3): 315-330.

[42] Gu J, Yin X, Struik P C, STOMPH T J, WANG H. Using chromosome introgression lines to map quantitative trait loci for photosynthesis parameters in rice (Oryza sativa L.) leaves under drought and well-watered field conditions. Journal of Experimental Botany, 2012, 63(1): 455-469.

[43] Griffiths H. Plant biology: designs on Rubisco. Nature, 2006, 441(7096): 940-941.