摘要
番茄(Lycopersicon esculentum Mill.)作为一种重要的经济作物,在世界范围内分布广泛。盐分胁迫通过破坏植物类囊体膜结构和功能导致光合性能下降,而类囊体膜上不饱和脂肪酸含量的上升则有利于光合功能的维持。番茄中决定叶绿体类囊体膜中磷脂酰甘油(PG)中顺式不饱和脂肪酸含量的是叶绿体甘油-3-磷酸酰基转移酶(LeGPAT)对底物的选择性。番茄中GPAT优先选择C18:1-ACP作为底物,因此番茄中PG的sn-1位上就含有较高比例的18:1脂肪酸,盐胁迫条件下,这些脂肪酸可以在酰基脂肪酸去饱和酶的作用下进一步去饱和化成为顺式多聚不饱和脂肪酸。
本文以野生型(WT),转正义番茄叶绿体甘油-3-磷酸酰基转移酶(LeGPAT)基因株系T2-19(+), T2-5(+)和转反义LeGPAT基因株系T2-16(-), T2-2(-)为试验材料,通过测定类囊体膜脂脂肪酸组成、盐胁迫后PG脂肪酸组成、光合及叶绿素荧光参数、叶绿体活性氧的产生、叶绿体活性氧清除酶活性以及D1蛋白修复等指标,研究了类囊体膜脂PG不饱和程度与番茄耐盐性的关系。主要结果如下:
1.对番茄植株类囊体膜脂脂肪酸组成进行分析,结果表明,在WT和转基因植株类囊体膜脂的四种类脂中双半乳糖基甘油二酯(DGDG),单半乳糖基甘油二酯(MGDG),硫代异鼠李糖基甘油二酯(SQDG)的脂肪酸含量没有明显变化,而PG中脂肪酸含量发生了明显变化。与野生型相比,转正义基因番茄植株类囊体膜脂PG中18:2和18:3含量明显增加,脂肪酸不饱和程度升高,而转反义基因番茄植株PG 18:2和18:3含量下降,脂肪酸不饱和度明显下降。盐胁迫下,转正义基因番茄植株PG不饱和脂肪酸含量比未处理前稍有增加,而WT和转反义基因株系不饱和脂肪酸含量稍有下降,总体变化不明显。
2.盐胁迫下,野生型和转基因株系的光合速率(Pn)和PSII最大光化学效率(Fv/Fm)都降低。但与野生型相比,转正义基因植株的Fv/Fm、Pn下降程度较小,而转反义基因植株的下降程度较大。表明类囊体膜脂PG不饱和度的提高有利于维持光合机构的稳定性,减轻对PSⅡ的光抑制。
3.与野生型相比,盐胁迫下转正义基因植株维持较高的叶绿体超氧化物歧化酶(SOD)和抗坏血酸过氧化物酶(APX)活性,产生较少的O2和H2O2;而转反义基因植株叶绿体抗氧化酶SOD及APX活性较低,O2和H2O2的含量较高。与其它株系相比,转正义基因株系叶绿体SOD和APX维持较高的活性,有利于清除活性氧,能够降低由于活性氧的积累而造成的膜脂过氧化程度,维持细胞膜的完整性。
4.分离类囊体膜,进行SDS-PAGE和Western杂交,分析盐胁迫对D1蛋白修复速率的影响。Western杂交表明,短期盐处理对D1蛋白含量影响较小。长期盐胁迫下野生型和转基因番茄植株D1蛋白总含量均下降。与野生型相比,转正义基因番茄植株能够维持较高的D1蛋白含量,转反义基因植株含量较低。D1蛋白合成抑制剂处理后,转正义基因番茄植株D1蛋白净降解速率较高;而转反义基因植株中D1蛋白净降解速率最慢。与野生型相比,盐胁迫下转正义基因番茄植株D1蛋白的修复速率最快,而转反义基因植株最慢。
上述结果表明,PG不饱和脂肪酸含量的增加可能提高D1蛋白的修复速率,提高叶绿体抗氧化酶的活性,进而提高番茄植株的耐盐性。
As an important cash crop, tomato (Lycopersicon esculentum mill.) is a world-wide distribution. Salt stress can damage the structure and function of thylakoid membrane, causing the decrease of photosynthesis, but the increase of unsaturated fatty acid of thylakoid membrane can enhance the ability of photosynthesis of plants. The dominant factor determining the level of cis-unsaturated fatty acids in PG is the substrate selectivity of glycerol-3-phosphate acyltransferase(LeGPAT)in tomato chloroplasts, which prefers 18:1-ACP to 16:0-ACP as a substrate. Therefore, a large proportion of oleic acid (18:1) occurs at the sn-1 position of PG in tomato, and under salt stress, oleic acid (18:1) of sn-1 position desaturates further into cis-polyunsaturated fatty acids of linoleic acid (18:2) and linolenic acid (18:3) by acyl-fatty acid desaturase in thylakoid membranes.
In order to investigate the relationship of fatty acid composition of PG in thylakoid membrane and the tolerance to salt stress, WT, sense transgenic lines T2-19(+), T2-5(+) and antisense transgenic lines T2-16(-), T2-2(-) of tomato were used to determine the fatty acid composition of thylakoid membrane, the fatty acid composition of PG, photosynthetic and chlorophyll fluorescence parameters, the production of chloroplastic reactive oxygen species (ROS) and activities of antioxidase in chloroplasts, and the repair of D1 under salt stress.
The results showed as follows:
1. The results showed that there was no significant difference about the unsaturated fatty acids composition of digalactosyldiacylglycerol (DGDG), monogalactosyldiacylglycerol (MGDG) and sulfoquinovosyldiacylglycerol (SQDG) in wild type (WT) and transgenic plants, but the content of fatty acids of PG changed markedly. Compared with WT, a higher content of 18:2 and 18:3 in PG was detected in sense transgenic plants. In contrast, the decreased unsaturation of PG in antisense transgenic plants was observed. And compared with WT, unsaturated fatty acids of PG increased in sense lines slightly, whereas decreased in WT and antisense ones lightly.
2. In order to investigate the correlation of fatty acid composition in thylakoid membrane and the tolerance of photosystem to salt stress, photosynthetic rate (Pn) and the maximal photochemical efficiency of PSII (Fv/Fm) in WT and transgenic plants decreased under salt stress, but compared with WT, Fv/Fm and Pn decreased more obviously in antisense lines, while more slowly in sense ones. It is indicated that the increase of unstaurated fatty acid in PG can stabilize the structure of photosynthetic apparatus, and alleviate the photoinhibition of PSII.
3. Compared with WT, the sense transgenic lines could maintain higher activities of chloroplastic SOD and APX, and produce lower content of O2 and H2O2 under salt stress. In contrast, the activities of chloroplastic SOD and APX were lower and the content of ROS was higher in the antisense transgenic plants. Relative to other lines, the higher activities of chloroplastic SOD and APX can scavenge more ROS, and mitigate the extent of lipid peroxidation caused by the accumulation of ROS, and maintain the integrity of the cytomembrane.
4. The analysis of western blot indicated that salt stress affected the repair of D1 protein. During short-time salt stress, the total content of D1 protein changed slightly among five lines. During salt stress for a long time, the content of D1 protein decreased in WT and transgenic plants. Compared with WT, there was a higher content of D1 protein in sense transgenic lines and lower level of D1 in antisense ones. After the treatment of streptomycin (SM), the net degradation of D1 was faster in sense trangenic plants and the most slowly in antisense ones. It is demonstrated that the rate of D1 turnover in sense lines was faster than other ones.
These results suggested that the increase of unsaturation in PG can enhance the salt tolerance in tomato plants by accelerating the repair of D1 protein and improving the activities of antioxidase in chloroplasts.
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