短时间热胁迫对疏叶骆驼刺光系统II、Rubisco活性和活性氧化剂的影响

doi:10.3724/SP.J.1258.2011.00441

植物生态学报 ›› 2011, Vol. 35 ›› Issue (4): 441-451.DOI: 10.3724/SP.J.1258.2011.00441

短时间热胁迫对疏叶骆驼刺光系统II、Rubisco活性和活性氧化剂的影响

薛伟¹^,²^,³^,⁴, 李向义¹^,³^,⁴^,^*(), 林丽莎¹^,³^,⁴, 王迎菊¹^,²^,³^,⁴, 李磊¹^,²^,³^,⁴

¹中国科学院新疆生态与地理研究所, 乌鲁木齐 830011
²中国科学院研究生院, 北京 100049
³新疆策勒荒漠草地生态系统国家野外科学观测研究站, 新疆策勒 848300
⁴中国科学院干旱区生物地理与生物资源重点实验室, 乌鲁木齐 830011

收稿日期:2010-11-05 接受日期:2011-01-21 出版日期:2011-11-05 发布日期:2011-04-13
通讯作者: 李向义
作者简介:*E-mail: lixy@ms.xjb.ac.cn

Effects of short time heat stress on photosystem II, Rubisco activities and oxidative radicals in Alhagi sparsifolia

XUE Wei¹^,²^,³^,⁴, LI Xiang-Yi¹^,³^,⁴^,^*(), LIN Li-Sha¹^,³^,⁴, WANG Ying-Ju¹^,²^,³^,⁴, LI Lei¹^,²^,³^,⁴

¹Xinjiang Institute of Ecology and Geography, Chinese Academy of Sciences, ürümqi 830011, China
²Graduate University of Chinese Academy of Sciences, Beijing 100049, China
³Cele National Station of Observation & Research for Desert Grassland Ecosystem, Cele, Xinjiang 848300, China
⁴Key Laboratory of Biogeography and Bioresource in Arid Land, Chinese Academy Sciences, ürümqi 830011, China

Received:2010-11-05 Accepted:2011-01-21 Online:2011-11-05 Published:2011-04-13
Contact: LI Xiang-Yi

摘要/Abstract

摘要：

生长在温带沙漠地区的植物在夏季时常遭受正午短时间的高温胁迫, 频繁和骤然的热胁迫在很大程度上限制了荒漠植物的光合作用。以塔克拉玛干沙漠南缘防风固沙的优势植物疏叶骆驼刺(Alhagi sparsifolia)为材料, 分别用叶绿素荧光诱导动力学和CO₂响应方法分析热胁迫后光系统II (PSII)和RuBP羧化酶的热稳定性。结果表明: (1)在叶片温度超过43 ℃后PSII最大光化学量子产量、有活性反应中心数目、活力指数均出现明显的降低; 中高温度下PSII的电子供体侧比电子受体侧组分更容易受到热胁迫的伤害; 在58 ℃出现明显的K点(300 μs), 说明放氧复合体放氧结构受到破坏而失去活性。(2)随着叶片温度的上升, Rubisco活性先升高后降低, 在34 ℃时具有最高的活性水平。(3)叶片受到高温胁迫时, 细胞内氨态氮和活性氧分子等大量积累。(4)疏叶骆驼刺叶片处于短时间的高温环境时, 光合作用的光反应和暗反应阶段均表现出功能的不稳定性, 其中PSII和Rubisco是主要的热敏感位点。

关键词: Rubisco活性, 疏叶骆驼刺, 热胁迫, 光系统II, 活性氧化剂

Abstract:

Aims Our general objective was to determine the effects of short time heat stress on photosystem II (PSII) activity and Rubisco activity in Alhagi sparsifolia. Specifically, we wanted to determine that temperature range within which the plant can photosynthesize, the critical temperature that damaged leaves and characters of reactive oxidative accumulation under high temperature.
Methods Samples of A. sparsifolia were collected on a sunny morning with low winds in early August 2010. We selected healthy growing shoots randomly, washed them lightly to remove leaf dust, cut and quickly inserted them into damp soil and covered them with a plastic cover. Samples were immediately put into black plastic bags, which contained damp filter papers. The bags were placed into a heat-resistant case for transport to the laboratory and then immersed in water baths for 15 min at temperatures of 30, 38, 43, 48, 53, 58 and 63 ℃. Twenty intact and mature leaves were used to measure leaf fluorescence and CO₂ response curves at each temperature.
Important findings At leaf temperatures up to 43 ℃, the maximum photochemical efficiency of PSII, number of active reaction centers and leaf vitality index all decreased markedly. At moderately high temperatures, the electron donors of PSII were more heat-vulnerable than the electron acceptors of PSII, and the appearance of ﬂuorescence K point (300 μs) in the ﬂuorescence curve at 58 ℃ indicated that the structure of oxygen-evolving complexes was damaged, resulting in loss of oxygen-evolving function. At higher leaf temperatures, the activity of Rubisco first increased and then decreased, with a maximum at 34 ℃. Under high temperature stress, considerable oxidative radicals, e.g., ammoniacal nitrogen, H₂O₂ and O₂^-· were produced and continually accumulated in cells. We conclude that heat stress has strong impacts on both light reaction and dark reaction phases of photosynthesis, especially their two heat sensitive components comprised of PSII and Rubisco.

Key words: activation of Rubisco, Alhagi sparsifolia, heat stress, photosystem II, oxidative radical

薛伟, 李向义, 林丽莎, 王迎菊, 李磊. 短时间热胁迫对疏叶骆驼刺光系统II、Rubisco活性和活性氧化剂的影响. 植物生态学报, 2011, 35(4): 441-451. DOI: 10.3724/SP.J.1258.2011.00441

XUE Wei, LI Xiang-Yi, LIN Li-Sha, WANG Ying-Ju, LI Lei. Effects of short time heat stress on photosystem II, Rubisco activities and oxidative radicals in Alhagi sparsifolia. Chinese Journal of Plant Ecology, 2011, 35(4): 441-451. DOI: 10.3724/SP.J.1258.2011.00441

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链接本文: https://www.plant-ecology.com/CN/10.3724/SP.J.1258.2011.00441

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图/表 8

图1 骆驼刺叶片Fv/Fm日变化(平均值±标准误差)。

Fig. 1 Daily changes of the maximum photochemical efficiency of PSII (Fv/Fm) of Alhagi sparsifolia leaves (mean ± SE).

图2 骆驼刺正常叶片典型的多相叶绿素荧光上升曲线O-J-I-P。各点的含义详见李鹏民等(2005)。

Fig. 2 A typical chlorophyll polyphasic ?uorescence rise O-J-I-P for an untreated Alhagi sparsifolia leaf. The meaning of each point referred Li et al. (2005).

图3 不同温度处理下骆驼刺叶片荧光动力学曲线的变化。

Fig. 3 Changes in chlorophyll ?uorescence induction curves in Alhagi sparsifolia leaves under different temperatures.

表1 热胁迫对叶绿素荧光参数的影响(平均值±标准误差)

Table 1 Effects of heat stress on chlorophyll ?uorescence parameters (mean ± SE)

处理 Treatment	初始荧光 F_o	最大荧光 F_m	可变荧光 F_v	PSII最大光化学量子产量 F_v/F_m
30 ℃	435.70 ± 21.04 (0%)	2 455.70 ± 147.46 (0%)	2 020.00 ± 139.72 (0%)	0.83 ± 0.01 (0%)
38 ℃	468.13 ± 24.13 (7.44%)	2 404.75 ± 134.34 (-2.07%)	1 936.63 ± 144.42 (-4.13%)	0.81 ± 0.02 (-2.41%)
43 ℃	464.30 ± 27.45 (6.56%)	2 318.40 ± 114.85 (-5.59%)	1 854.10 ± 153.18 (-8.21%)	0.80 ± 0.01 (-3.61%)
48 ℃	602.00 ± 33.20 (38.17%)	1 948.40 ± 103.69 (-20.66%)	1 346.40 ± 62.77 (-33.35%)	0.69 ± 0.06 (-16.87%)
53 ℃	604.13 ± 33.42 (38.66%)	1 840.75 ± 89.03 (-25.04%)	1 236.63 ± 94.79 (-38.78%)	0.65 ± 0.10 (-21.69%)
58 ℃	849.40 ± 38.11 (94.95%)	1 433.80 ± 69.32 (-41.61%)	584.40 ± 25.67 (-71.07%)	0.41 ± 0.10 (-50.60%)
63 ℃	1 807.80 ± 40.02 (314.92%)	1 809.20 ± 106.7 (-26.33%)	1.40 ± 1.04 (-99.93%)	0.000 7 ± 0.000 1 (-99.92%)

图4 热胁迫对骆驼刺叶片PSII能量流动(A)和能量利用效率(B)的影响(平均值±标准误差)。图中曲线的颜色与纵坐标颜色相对应。ABS/RC, 天线色素吸收的能量; DIo/RC, 用于热耗散的能量; ETo/RC, 用于电子传递的能量; Fv/Fm, PSII最大光化学量子产量; PIABS, 叶片性能指数; RC/CS, 单位面积的光合机构被激活反应中心的数量; TRo/RC, 反应中心捕获的能量; ρk, 放氧复合体放氧活性; ΦEo, 电子传递的量子产额; ψo, 捕获的激发能导致电子传递的效率。

Fig. 4 Effects of heat stress on PSII energy flux (A) and energy use efficiency (B) of Alhagi sparsifolia leaves (mean ± SE). In graph the color of curve is correspondence with the color of y-axis. The specific energy fluxes (per reaction centers, RC) for absorption (ABS/RC), trapping (TRo/RC), electron transport (ETo/RC) and dissipation (DIo/RC); the flux ratios or yield, i.e. the maximum quantum yield of primary photochemistry (Fv/Fm), the efficiency with which a trapped exciton can move an electron into the electron transport chain further than OA- (ψo), and the quantum yield of electron transport (ΦEo); the fraction of O2 evolving centers in comparison with the control sample (ρk); the amount of active PSII reaction centers per excited cross section (RC/CS), and the performance index (PIABS).

图5 高温胁迫对骆驼刺叶片RuBP羧化效率的影响(平均值±标准误差)。计算初始斜率时采用前6个点进行拟合。

Fig. 5 Effects of elevated temperature on RuBP carboxylation efficiency of Alhagi sparsifolia leaves (mean ± SE). Parameters of linear fitting based on the initial stage of Pn-Ci cruves which only contains the six foremost data points.

图6 热胁迫对疏叶骆驼刺叶片中氨态氮(A)、活性氧分子O2-· (B)和H2O2 (C)的影响(平均值±标准误差)。

Fig. 6 Effects of heat stress on ammoniacal nitrogen (A) and reactive oxygen molecules O2-· (B) and H2O2 (C) concentration of Alhagi sparsifolia leaves (mean ± SE).

图7 光合系统主要的热胁迫伤害位点(绘制结构参考于http://i.hudong.com/profile.do?useriden=nfEACAwQFWUICDQJ_和http://tupian.hudong.com/a3_42_21_01300000332400124108219709521_jpg.html。但光合反应的生理学过程经过了本文作者思考加工)。2,3-DPG, 二磷酸甘油酸; Cyt.b6, 细胞色素b6-f复合物; Fd, 铁氧还原蛋白; LHCII, 光系统II捕光色素系统; OEC, 放氧复合体; P680, 光系统II反应中心色素; P700, 光系统I反应中心色素; PC, 质体蓝素; PGA, 磷酸甘油酸; Pheo, 去镁叶绿素; PQ, 质体醌; QA, 初级醌受体; QB, 次级醌受体; Sucrose, 蔗糖; Tyiose-P, 磷酸丙糖; Yz, 原初电子供体。

Fig. 7 The heat damaged parts of photosynthesis which includes light reaction phase mainly PSII and dark reaction phase under high temperature. When green leaves were exposed to high temperature (e.g. over 43 ℃), three important components given above oxygen evolving complex (OEC), light-harvesting system and Rubisco were more vulnerable than others components. 2,3-DPG, 2, 3-diphosphoglycerate; Cyt.b6, cytochrome b6-f complex; Fd, ferredoxin; LHCII, light-harvesting complex in photosystem II; P680, PSII reaction centre; P700, photosystem I reaction centre; PC, plastocyanin; PGA, phosphoglycerate; Pheo, pheophytin; PQ, plastoquinone; QA, the primary quinone acceptor; QB, the secondary quinone acceptor; Tyiose-P, triose phosphate; Yz, tyrosine Z.

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短时间热胁迫对疏叶骆驼刺光系统II、Rubisco活性和活性氧化剂的影响

Effects of short time heat stress on photosystem II, Rubisco activities and oxidative radicals in Alhagi sparsifolia

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