青藏高原春小麦叶片光合作用的光抑制及PSII反应中心光化学效率的恢复分析

doi:10.3724/SP.J.1258.2014.00034

摘要/Abstract

摘要：

在青海省都兰县香日德镇东盛村, 以中国科学院西北高原生物研究所培育的春小麦(Triticum aestivum)品种为材料, 主要采用调制叶绿素荧光分析手段, 研究了抽穗期旗叶光合作用的光抑制现象, 并分析了非光化学猝灭组分的光诱导和非光诱导耗散的量子产量变化。结果表明, 高原春小麦各品种间旗叶光合色素含量和比叶重存在差异; 全晴天3个典型时段准确暗适应20 min后的PSII最大光化学效率(F_v/F_m)的比较分析证实, 高原春小麦存在着光合作用的光抑制现象, F_v/F_m的降低是由于PSII反应中心的可逆失活; 稳态作用光下PSII有效光化学效率(F_v′/F_m′)易受持续强光胁迫的影响, 而PSII实际光化学效率(Φ_PSII)在各春小麦品种间的差异略为明显; 上下午间4个春小麦品种的光化学猝灭系数(q_P)和非光化学猝灭系数(NPQ)呈较一致的变化趋势, 显然q_P和NPQ既属品种的内禀特性, 又与强太阳光胁迫的累积密切相关; 非光化学猝灭组分中光诱导的PSII调节性能量耗散的量子产量(Φ_NPQ)所占比例较大, 下午时分Φ_NPQ的上调反映了高原春小麦对青藏高原持续强光胁迫的驯化适应。

关键词: 叶绿素荧光, 强光胁迫, 光抑制, PSII光化学效率, 青藏高原, 春小麦

Abstract:

Aims Several local varieties of wheat (Triticum aestivum) developed by Northwest Institute of Plateau Biology, Chinese Academy of Sciences, are widely cultivated in the agricultural regions in Qinghai-Xizang Plateau. These varieties are well adapted to multiple environmental stresses such as low temperature, strong solar radiation, and drought. The objective of this study was to determine the responses of PSII photochemical efficiency to high solar irradiance in leaves of four wheat varieties. We examined whether photo-inhibition was appeared in wheat varieties and analysed variations of quantum yield of quenching due to light-induced and non-light-induced.
Methods Field experiments were conducted on the farmland of Xiangride, which is located in the eastern side of Caidamu Basin, Qinghai Province. Four local wheat varieties were used during the heading stage in 2013. Measurements of photochemical efficiency and quantum yield were made on the abaxial surface of flag leaves facing the Sun by using a FMS-2 fluorometer, and the content of photosynthetic pigments and specific leaf weight (SLW) were concurrently determined. Pulse-modulated in-vivo chlorophyll fluorescence technique was used to obtain rapid information on photosynthetic processes. The maximum quantum efficiency of PSII photochemistry (F_v/F_m) was determined at 8:30, 12:00 and 16:30 on clear days after allowing for 20 min dark adaptation with leaf clips. The PSII maximal and actual photochemical efficiency (F_v′/F_m′ and Φ_PSII), the PSII photochemical and non-photochemical quenching coefficient (q_P and NPQ) were analyzed between morning and afternoon using inner actinic light with photosynthetically active photon flux density at 1120 μmol photons∙m^-2∙s^-1. Furthermore, along with analysis of the fraction of PSII reaction centers that are opened (q_L), the quantum yield of quenching due to light-induced processes (Φ_NPQ) and non-light-induced processes (Φ_NO) were explored.
Important findings There were significant differences in the content of photosynthetic pigments and SLW among the four wheat varieties. Under conditions of clear days, the flag leaves exhibited marked depressions in F_v/F_m at three typical times when determined after 20 min dark adaptation. At a given light intensity, the values of F_v′/F_m′ were significantly reduced in the afternoon due to influences by long-lasting high-light irradiation, and Φ_PSII showed little differences among the four wheat varieties and no difference between morning and afternoon. There were almost similar variations in q_P and NPQ among the four wheat varieties, suggesting that q_P and NPQ belong to instinct property and are influenced by the accumulative stresses of high-light intensity. The fractions of Φ_NPQ were higher than that of Φ_NO in the four wheat varieties and the up-regulatory of Φ_NPQ in the afternoon indicated that the photosynthetic apparatus in these wheat varieties had already acclimated to strong solar irradiation in agricultural regions of Qinghai-Xizang Plateau.

Key words: chlorophyll fluorescence, high-light stress, photosynthetic photo-inhibition, PSII photochemical efficiency, Qinghai- Xizang Plateau, spring wheat

师生波, 张怀刚, 师瑞, 李妙, 陈文杰, 孙亚男. 青藏高原春小麦叶片光合作用的光抑制及PSII反应中心光化学效率的恢复分析. 植物生态学报, 2014, 38(4): 375-386. DOI: 10.3724/SP.J.1258.2014.00034

SHI Sheng-Bo, ZHANG Huai-Gang, SHI Rui, LI Miao, CHEN Wen-Jie, SUN Ya-Nan. Assessment of photosynthetic photo-inhibition and recovery of PSII photochemical efficiency in leaves of wheat varieties in Qinghai-Xizang Plateau. Chinese Journal of Plant Ecology, 2014, 38(4): 375-386. DOI: 10.3724/SP.J.1258.2014.00034

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https://www.plant-ecology.com/CN/Y2014/V38/I4/375

图/表 7

图1 抽穗期4个高原春小麦品种旗叶的光合色素含量的变化(平均值±标准偏差, n = 6)。不同小写字母表示春小麦品种间差异显著(p = 0.05)。

Fig. 1 Variations in the content of photosynthetic pigments in the flag leaves of four wheat varieties during the heading stage (mean ± SD, n = 6). Different lower-case letters indicate significant differences among wheat varieties (p = 0.05).

图2 抽穗期4个高原春小麦品种旗叶的比叶干重(SLWd) (A)和比叶鲜重(SLWf) (B)的变化(平均值±标准偏差, n = 15)。不同小写字母表示春小麦品种间的差异显著(p = 0.05)。

Fig. 2 Variations in the specific leaf dry weight (SLWd) (A) and specific leaf fresh weight (SLWf) (B) in the flag leaves of four wheat varieties during the heading stage (mean ± SD, n = 15). Different lower-case letters indicate significant differences among wheat varieties (p = 0.05).

图3 抽穗期全晴天3个典型时间段20分钟暗适应后4个高原春小麦品种旗叶的PSII最大光化学量子效率(Fv/Fm和1/Fo - 1/Fm)的变化(平均值±标准偏差, n = 12)。不同小写字母表示4个春小麦品种在全晴天3个时间段之间的差异显著(p = 0.05)。

Fig. 3 Variations in the maximum photochemical efficiency of PSII (Fv/Fm and 1/Fo - 1/Fm) in the flag leaves of four wheat varieties after 20 min dark adaptation at three measurement times on a clear day during the heading stage (mean ± SD, n = 12). Different lower-case letters indicate significant differences among three typical times within wheat varieties on a clear day (p = 0.05).

图4 抽穗期全晴天3个典型时间段4个高原春小麦品种旗叶PSII反应中心叶绿素初始荧光产量(Fo)的变化(平均值±标准偏差, n = 12)。不同小写字母表示4个春小麦品种在3个时间段之间的差异显著(p = 0.05)。

Fig. 4 Variations in the minimal fluorescence of PSII reaction centers (Fo) in the flag leaves of four wheat varieties after 20 min dark adaptation at three typical times during the heading stage (mean ± SD, n = 12). Different lower-case letters in each column indicate significant differences among three typical times within wheat varieties on a clear day (p = 0.05).

图5 抽穗期晴天稳定作用光下4个春小麦品种旗叶的PSII反应中心有效光化学量子效率(Fv′/Fm′)(A)和实际光化学量子效率(ΦPSII)(B)及上下午差异分析(平均值±标准偏差, n = 6)。不同大写字母和小写字母分别表示上午和下午4个春小麦品种旗叶的Fv′/Fm′和ΦPSII的差异显著(p = 0.05)。ns, 上下午间无显著差异(p > 0.05); *和**, 上下午间差异显著和极显著(p < 0.05, p < 0.01)。

Fig. 5 Analysis of the PSII maximal photochemical efficiency (Fv′/Fm′) (A) and actual photochemical efficiency (ΦPSII) (B) in the flag leaves of four wheat varieties between morning and afternoon at a given light intensity during the heading stage (mean ± SD, n = 6). Different capital and lower-case letters in figures indicate significant differences in Fv′/Fm′ and ΦPSII, respectively, between morning and afternoon (p = 0.05). ns, no significant differences between morning and afternoon (p > 0.05); * and **, significant and highly significant differences between morning and afternoon (p < 0.05 and p < 0.01).

图6 抽穗期晴天稳定作用光下4个春小麦品种旗叶的PSII反应中心光化学猝灭系数(qP)(A)和非光化学猝灭系数(NPQ) (B)及上下午差异分析(平均值±标准偏差, n = 6)。不同大写字母和小写字母分别表示上午和下午间4个春小麦品种旗叶的qP和NPQ的差异显著(p = 0.05)。ns, 上下午间无显著性差异(p > 0.05); *和**, 上下午间差异显著和极显著(p < 0.05和 p < 0.01)。

Fig. 6 Analysis of PSII photochemical quenching coefficient (qP) (A) and non-photochemical quenching coefficient (NPQ) (B) in the flag leaves of four wheat varieties between morning and afternoon at a given light intensity during the heading stage (means ± SD, n = 6). Different capital and lower-case letters in figures indicate significant differences in qP and NPQ, respectively, between morning and afternoon (p = 0.05). ns, no significant differences between morning and afternoon; * and **, significant and highly significant differences between morning and afternoon (p < 0.05 and p < 0.01).

图7 抽穗期高原春小麦品种旗叶PSII反应中心开放比率(qL)(A)及调节性能量耗散量子产量(ΦNPQ)(B)和非调节性能量耗散量子产量(ΦNO)(C)的上下午的变化(平均值±标准偏差, n = 6)。不同大写字母和小写字母分别为上午和下午4个春小麦品种旗叶的qL、ΦNPQ和ΦNO差异显著(p = 0.05)。ns, 上下午间无显著性差异(p > 0.05); *和**, 上下午间差异显著和极显著(p < 0.05和p < 0.01)。

Fig. 7 Variations in the fraction of PSII reaction centers that are open (qL) (A), PSII regulatory energy dissipation in quantum yield (ΦNPQ) (B), and non-regulatory energy dissipation in quantum yield (ΦNO) (C) in the flag leaves of four wheat varieties between morning and afternoon at the given light intensity during the heading stage (means ± SD, n = 6). Different capital and lower-case letters in figures indicate significant differences in qL, ΦNPQ and ΦNO, respectively, between morning and afternoon (p = 0.05). ns, no significant differences between morning and afternoon; * and **, significant and highly significant differences between morning and afternoon (p < 0.05 and p < 0.01).

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