碱胁迫对小麦（Triticum aestivum Linn）叶片代谢过程的影响

doi:10.3864/j.issn.0578-1752.2017.02.005

摘要/Abstract

摘要： 【目的】阐明碱胁迫对小麦叶片离子平衡、初生及次生代谢产物的影响及其涉及的代谢途径，讨论其生长代谢变化规律及应答机制。【方法】以普通小麦（Triticum aestivum Linn）为材料，采用盆栽试验利用NaHCO₃﹕Na₂CO₃=1﹕1混合模拟不同盐度碱胁迫条件，在苗期连续胁迫12 d后测定叶片生长、光合、离子和代谢产物。【结果】当碱胁迫强度超过小麦自身调节能力时，叶片中Na⁺含量剧增，加上高pH危害，造成叶绿体遭到破坏、叶绿素含量降低、光系统Ⅱ活性受抑制、气孔导度及碳同化能力急剧下降，最终导致生长率降低。碱胁迫下Na⁺大量增加的同时阴离子明显减少，造成叶片内负电荷亏缺和pH不稳定，导致离子平衡遭到破坏，进而引起一系列代谢途径的协变反应。通过GC-MS检测出73个代谢物，主要包括碳水化合物、氨基酸、有机酸等，其中，分别有25和48个代谢物在中度和重度碱胁迫下发生明显改变。主成分分析（PCA）结果显示全部样本均分布在95%的置信区间内，2个主成分得分达到89%。单因素方差分析表明，与对照组比较，在高浓度碱胁迫下发生的显著性变化明显高于低浓度碱胁迫。碱胁迫导致5种参与三羧酸（TCA）循环和6种参与糖酵解途径的代谢物含量明显降低，且引起大部分氨基酸（谷氨酸、丙氨酸、γ-氨基丁酸、天冬氨酸等）和糖类及多元醇（果糖、蔗糖、塔罗糖、肌醇等）大量降低。与此同时，碱胁迫诱导小麦有机酸大量积累，随胁迫强度的增加而上升，这种现象可能是小麦被动的适应调节过程，主要用于维持离子平衡并调节pH浓度。【结论】碱胁迫引起了TCA循环、糖酵解途径、卡尔文循环、莽草酸途径、细胞膜脂代谢、转氨基反应和γ-氨基丁酸（GABA）途径等代谢网络系统广泛变化，暗示了碱胁迫不仅对糖类、氨基酸类、脂肪和蛋白质合成代谢过程造成负面影响，而且限制C-N转变过程影响植物对N素的利用，造成营养匮乏抑制植物生长发育。

关键词: 小麦, 碱胁迫, 叶片, 生长特性, 光合特性, 代谢组

Abstract: 【Objective】 A pot experiment was conducted to investigate the alkaline stress in ion balance and metabolic profiles of wheat, to determine the physiological adaptive mechanisms of wheat in tolerance to alkali stress. 【Method】 In a pot experiment with control and alkaline stress (NaHCO₃﹕Na₂CO₃=1﹕1), the growth and photosynthetic characters, ion and 73 key metabolites of wheat were studied. 【Result】The results showed that when alkaline stress intensity exceeded the capacity of wheat adjustment, Na⁺ accumulation in cells in a high-pH environment resulted in damage of the photosynthetic system, reduced photosynthetic pigments, inhibited the activity of photosystem II, and reduced high stomatal conductance and net photosynthetic rate. Alkaline stress caused massive influx of Na⁺, a decrease of inorganic negative charge and pH value homeostasis, thus resulting ionic unbalance and leading to a series of strain metabolic response. In addition, 73 metabolites were detected in different alkaline stress treatments according to GC-MS analysis, and these metabolites were sugars/polyols, organic acids, amino acids and others. Compared with the control sample, the response of 25 and 48 metabolites in moderate and severe alkaline stress treatments remarkably changed, respectively, in leaves of wheat seedlings. The results of one-way ANOVA analysis indicated that the changes of metabolites were more significant under high alkaline stress than that under moderate alkaline stress. The results revealed that alkaline stress caused an significant decrease in levels of 5 and 6 metabolites, which are involved in TCA cycle and glycolysis; it also caused amino acids (glutamate, alanine, γ-aminobutyric acid, aspartic acid) and sugars/ploys (fructose, sucrose, talose, myo-inositol) decreased dramatically. Meanwhile, alkaline stress induced organic acids accumulation in wheat, and it maybe a passive adaptive response to alkaline stress, and organic acids kept ionic balance and pH homeostasis. 【Conclusion】 The results suggested that alkaline stress caused systems alterations in metabolic networks including TCA cycle, glycolysis, calvin cycle, shikimic path way, metabolism of plasma membrane,GS/GOGAT cycle and GABA path way, implying alkaline stress not only had side effect on synthesis of sugars, amino acids, fats and proteins, but also inhibited the translation between C and N, thus resulted in nutrients deficiency and caused decrease of plant growth and development.

Key words: wheat (Triticum aestivum Linn), alkali stress, leaves, growth characters, photosynthetic characters, metabonomic

郭瑞，周际，杨帆，李峰. 碱胁迫对小麦（Triticum aestivum Linn）叶片代谢过程的影响[J]. 中国农业科学, 2017, 50(2): 250-259.

GUO Rui, ZHOU Ji, YANG Fan, LI Feng. Effects of Alkaline Stress on Metabonomic Responses of Wheat (Triticum aestivum Linn) Leaves[J]. Scientia Agricultura Sinica, 2017, 50(2): 250-259.

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