铝诱导油菜有机酸分泌和代谢的调控及铝毒特性研究
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摘要
Al胁迫下植物根系大量分泌有机酸的可能调节因素主要包括两个方面,一是有机酸代谢:Al诱导根系细胞内有机酸合成量的增加而使其大量积累而最终向外分泌;二是有机酸运输:Al选择性地诱导开启细胞膜上特定有机酸阴离子通道使有机酸的大量分泌,线粒体膜上的载体也可能是调节有机酸向外分泌的关键因素。在前人的研究基础上,本文以有机酸分泌类型属于模式Ⅱ植物的油菜作为研究对象,通过在AlCl_3胁迫下添加外源阴离子通道抑制剂A9C和柠檬酸载体抑制剂Mersal,比较两者不同的抑制剂对于铝毒诱导油菜根系分泌有机酸的调控,以及油菜体内的有机酸代谢对添加有抑制剂的铝毒胁迫的响应,进一步探讨有机酸分泌与有机酸代谢之间的关系;此外,还从生长生理和营养吸收等方面阐述外源抑制剂对油菜Al毒毒害特征的影响。结果表明:
1.在200μmol/L AlCl_3胁迫下,油菜根系分泌的柠檬酸比对照增加3.4倍,苹果酸比对照增加4.4倍,柠檬酸和苹果酸占据有机酸分泌总量的70%以上。A9C和Mersal对Al诱导的油菜根系有机酸分泌具有抑制作用。A9C对苹果酸分泌的抑制较大,5μmol/L和20μmol/L浓度下苹果酸分泌比单Al处理下分别下降15%和35%;Mersal对柠檬酸分泌的抑制较大,5μmol/L和20μmol/L浓度下柠檬酸分泌分别下降13%和28%。由此说明,油菜根系中的阴离子通道和柠檬酸载体均对有机酸分泌具有调节作用,并且两者具有联动效应。
2.在200μmol/L AlCl_3胁迫下,油菜根系和叶片中的柠檬酸和苹果酸合成也比对照显著增加,柠檬酸分别增加了111%和103%,苹果酸分别增加了223%和147%。A9C和Mersal同样对铝毒胁迫下油菜体内有机酸的合成具有抑制作用,但仍然显著高于对照。实验还发现,柠檬酸和苹果酸在根中的合成要大于在叶中的合成。相关性分析表明,柠檬酸和苹果酸的分泌与根系和叶片内柠檬酸和苹果酸的合成均达到显著相关,且根系中的有机酸和叶片中的有机酸也达到极显著相关。说明调控有机酸分泌的阴离子通道和柠檬酸载体也影响油菜体内有机酸的合成。
3.油菜在受到AlCl_3胁迫后,根系和叶片中的CS、MDH、PEPC、ACO活性显著高于对照,前三者增加率均大于100%,而ACO活性增加较小,这样的变化趋势有利于柠檬酸和苹果酸在根系和叶片中的积累。进行抑制剂-Al复合处理之后,与单Al处理比较,油菜根系和叶片中的CS、MDH和PEPC活性均有不同程度的下降,而ACO活性相反却有所升高。结果说明,抑制剂-Al毒胁迫导致了油菜组织中柠檬酸和苹果酸的合成能力减弱,促进了柠檬酸向乌头酸的转变,进一步降低了组织中的柠檬酸含量,并最终减少了油菜有机酸的分泌。
4.Al胁迫使油菜根系分泌氨基酸和可溶性糖含量增加,但与对照差异不显著;抑制剂-Al复合处理后,氨基酸和可溶性糖的渗透增强,20μmol/L的A9C使氨基酸和可溶性糖的渗透明显增多,与对照和单Al处理差异显著(P＜0.05)。根质膜透性在单Al胁迫下比对照有所升高,A9C和Mersal加剧了铝对细胞质膜的损伤,使根质膜透性上升到与对照差异显著。研究认为,A9C和Mersal导致Al胁迫下氨基酸和可溶性糖渗透增加是细胞质膜严重损伤的结果。单Al处理下根系分泌物的pH值比对照增加,认为是油菜对铝毒的适应性反应;A9C和Mersal加入导致了pH值比对照有所下降,可能是两种抑制剂加剧油菜Al毒害引起。
5.在铝液中添加A9C和Mersal后,油菜根系、地上部干重及根/冠比进一步下降,主根长、株高、叶片数增长率明显受到抑制,根系各形态指标(总根长、总表面积、平均直径、总体积、根尖数)下降增加,根系和地上部含水率下降。此外,油菜根、茎、叶中的N、P、K、Ca、Mg含量均比单Al处理下有不同程度的下降,Na含量却有所上升,从而提高了油菜根、茎、叶中的Na~+/K~+;并且,A9C和Mersal均促进了铝毒胁迫下油菜根、茎、叶对Al的吸收。结果说明添加A9C和Mersal可能减少了油菜体内合成和根际分泌的有机酸,从而降低了有机酸与Al的络合作用而使铝毒伤害加深。
The efflux of organic acid anions from plant roots under aluminum (Al) were likely related to two aspects, one was metabolism of organic acids, which Al increased the synthesis of organic acid in root-cells to increase accumulation of organic acid in plant, finally induced organic acid excreted; the other was transport of organic acids, which Al induced the opening of organic acid anion channel to induced organic acid excreted, and the carrier on the mitochondrial membrane was also may be the key regulation factor of organic acid excreted. Bases of the previously research, oilseed rape which belonged to patternⅡof exudation of organic acid was taken as object for the study, by the way of supplying A9C (a inhibitor of anion channel) and Mersal (a inhibitor of citrate carrier) in the Al~(3+) solution, the purpose of the study was to compare the difference of two inhibitor on the regulation of organic acid excreted, and the response of metabolism of organic acid to Al stress which contained inhibitor, thus explored the relation between metabolism and excretion of organic acid. Besides, the physiological characteristics and mineral nutrient absorb of oilseed rape under Al stress were investigated in the study. The main results were summarized as follows:
Under 200μmol/L AlCl_3 stress, the secretion of citric acid and malic acid increased respectively 3.4 times and 4.4 times compared to CK, at the time both of the two organic acids took up more than 70% in total organic acids secretion. A9C and Mersal could inhibit the secretion of organic acid which was induced by Al, compared to single Al-treatment, A9C inhibited significantly the malic acid by 15% and 35% under 5μmol/L and 20μmol/L of concentration, the corresponding inhibition for Mersal to citric acid were 13% and 28%. These results indicated that both of anion channel and citrate carrier could regulated the secretion of organic acid, and it had association between the two.
Under 200μmol/L AlCl_3 stress, the synthesis of citric acid and malic acid in roots and leaves increased too compared to CK, which citric acid increased by 111% and 103%, malic acid increased by 223% and 147%, respectively. As same as excretion of organic acid, A9C and Mersal could inhibit the synthesis of organic acid which was induced by Al, but that higher markedly than CK. It was found that the synthesis of organic acid in roots was intensive that of in leaves. The excretion of organic acid and the synthesis of citric acid and malic acid in roots or leaves was a significant positive correlation. These indicated that anion channel and citrate carrier not only regulated excretion of organic acid, but also affected synthesis of organic acid.
The activities of CS, MDH, PEPC and ACO of roots and leaves increased significantly in oilseed rape under single Al stress, the increasing ratios of CS, MDH, PEPC were over 100%, whereas ACO increased less than the three. The trend of change made for accumulation of citric acid and malic acid in roots and leaves. With adding inhibitors into AlCl_3 culture solution, the activities of CS, MDH, PEPC decreased with different degree, ACO was reverse, compared to single Al treatment. These indicated that inhibitor-Al treatments depressed the ability of synthesis of organic acid in plant, and may to citric acid transformed into aconitic acid, thus reduced the contents of citric acid in organization of plant, finally decreased the excretion of organic acid.
The contents of amino acid and soluble sugar increased under Al stress, but it was not significantly difference compared to CK. Complex treatments more increased the osmosis of amino acid and soluble sugar in the root exudation, especially 20μmol/L of A9C that reached markedly level (P<0.05) compared to single Al treatment and CK. The permeability of plasma membrane was increased under Al stress, especially added the A9C and Mersal in the Al solution. It was considered that the osmosis increasing of amino acid and soluble sugar was result of the damage of plasma membrane damage by Al stress. The increasing of pH in excretion under single Al stress was considered an adaptive response on Al stress, and the pH declining after subjoining A9C and Mersal in Al solutions was result in more toxicity of Al to oilseed rape seedlings.
When addition of A9C or Mersal under the Al stress, the dry weights of root and shoot and the ratio of root/shoot decreased rapidly, the growth ratios of main root, plant height and leaf amount inhibited significantly, the root morphology (root length, surface area, average diameter of root, root volume and number of root tips) and moisture contents of root and shoot descended too. In addition, the contents of N, P, K, Ca and Mg in root, stem and leaf of plant declined while Na increased in different extent compared to single Al treatment, which consequently enhanced the ratio of Na~+/ K~+ of root, stem and leaf. Moreover, the accumulation of Al in root, stem and leaf increased obviously with adding A9C and Mersal. It was suggested that A9C and Mersal might decrease the synthesis or excretion of organic acid in oilseed rape, therefore reduced the combine to organic acid with Al, finally the Al toxicity was fierce in plant.
1.在200μmol/L AlCl_3胁迫下,油菜根系分泌的柠檬酸比对照增加3.4倍,苹果酸比对照增加4.4倍,柠檬酸和苹果酸占据有机酸分泌总量的70%以上。A9C和Mersal对Al诱导的油菜根系有机酸分泌具有抑制作用。A9C对苹果酸分泌的抑制较大,5μmol/L和20μmol/L浓度下苹果酸分泌比单Al处理下分别下降15%和35%;Mersal对柠檬酸分泌的抑制较大,5μmol/L和20μmol/L浓度下柠檬酸分泌分别下降13%和28%。由此说明,油菜根系中的阴离子通道和柠檬酸载体均对有机酸分泌具有调节作用,并且两者具有联动效应。
2.在200μmol/L AlCl_3胁迫下,油菜根系和叶片中的柠檬酸和苹果酸合成也比对照显著增加,柠檬酸分别增加了111%和103%,苹果酸分别增加了223%和147%。A9C和Mersal同样对铝毒胁迫下油菜体内有机酸的合成具有抑制作用,但仍然显著高于对照。实验还发现,柠檬酸和苹果酸在根中的合成要大于在叶中的合成。相关性分析表明,柠檬酸和苹果酸的分泌与根系和叶片内柠檬酸和苹果酸的合成均达到显著相关,且根系中的有机酸和叶片中的有机酸也达到极显著相关。说明调控有机酸分泌的阴离子通道和柠檬酸载体也影响油菜体内有机酸的合成。
3.油菜在受到AlCl_3胁迫后,根系和叶片中的CS、MDH、PEPC、ACO活性显著高于对照,前三者增加率均大于100%,而ACO活性增加较小,这样的变化趋势有利于柠檬酸和苹果酸在根系和叶片中的积累。进行抑制剂-Al复合处理之后,与单Al处理比较,油菜根系和叶片中的CS、MDH和PEPC活性均有不同程度的下降,而ACO活性相反却有所升高。结果说明,抑制剂-Al毒胁迫导致了油菜组织中柠檬酸和苹果酸的合成能力减弱,促进了柠檬酸向乌头酸的转变,进一步降低了组织中的柠檬酸含量,并最终减少了油菜有机酸的分泌。
4.Al胁迫使油菜根系分泌氨基酸和可溶性糖含量增加,但与对照差异不显著;抑制剂-Al复合处理后,氨基酸和可溶性糖的渗透增强,20μmol/L的A9C使氨基酸和可溶性糖的渗透明显增多,与对照和单Al处理差异显著(P＜0.05)。根质膜透性在单Al胁迫下比对照有所升高,A9C和Mersal加剧了铝对细胞质膜的损伤,使根质膜透性上升到与对照差异显著。研究认为,A9C和Mersal导致Al胁迫下氨基酸和可溶性糖渗透增加是细胞质膜严重损伤的结果。单Al处理下根系分泌物的pH值比对照增加,认为是油菜对铝毒的适应性反应;A9C和Mersal加入导致了pH值比对照有所下降,可能是两种抑制剂加剧油菜Al毒害引起。
5.在铝液中添加A9C和Mersal后,油菜根系、地上部干重及根/冠比进一步下降,主根长、株高、叶片数增长率明显受到抑制,根系各形态指标(总根长、总表面积、平均直径、总体积、根尖数)下降增加,根系和地上部含水率下降。此外,油菜根、茎、叶中的N、P、K、Ca、Mg含量均比单Al处理下有不同程度的下降,Na含量却有所上升,从而提高了油菜根、茎、叶中的Na~+/K~+;并且,A9C和Mersal均促进了铝毒胁迫下油菜根、茎、叶对Al的吸收。结果说明添加A9C和Mersal可能减少了油菜体内合成和根际分泌的有机酸,从而降低了有机酸与Al的络合作用而使铝毒伤害加深。
The efflux of organic acid anions from plant roots under aluminum (Al) were likely related to two aspects, one was metabolism of organic acids, which Al increased the synthesis of organic acid in root-cells to increase accumulation of organic acid in plant, finally induced organic acid excreted; the other was transport of organic acids, which Al induced the opening of organic acid anion channel to induced organic acid excreted, and the carrier on the mitochondrial membrane was also may be the key regulation factor of organic acid excreted. Bases of the previously research, oilseed rape which belonged to patternⅡof exudation of organic acid was taken as object for the study, by the way of supplying A9C (a inhibitor of anion channel) and Mersal (a inhibitor of citrate carrier) in the Al~(3+) solution, the purpose of the study was to compare the difference of two inhibitor on the regulation of organic acid excreted, and the response of metabolism of organic acid to Al stress which contained inhibitor, thus explored the relation between metabolism and excretion of organic acid. Besides, the physiological characteristics and mineral nutrient absorb of oilseed rape under Al stress were investigated in the study. The main results were summarized as follows:
Under 200μmol/L AlCl_3 stress, the secretion of citric acid and malic acid increased respectively 3.4 times and 4.4 times compared to CK, at the time both of the two organic acids took up more than 70% in total organic acids secretion. A9C and Mersal could inhibit the secretion of organic acid which was induced by Al, compared to single Al-treatment, A9C inhibited significantly the malic acid by 15% and 35% under 5μmol/L and 20μmol/L of concentration, the corresponding inhibition for Mersal to citric acid were 13% and 28%. These results indicated that both of anion channel and citrate carrier could regulated the secretion of organic acid, and it had association between the two.
Under 200μmol/L AlCl_3 stress, the synthesis of citric acid and malic acid in roots and leaves increased too compared to CK, which citric acid increased by 111% and 103%, malic acid increased by 223% and 147%, respectively. As same as excretion of organic acid, A9C and Mersal could inhibit the synthesis of organic acid which was induced by Al, but that higher markedly than CK. It was found that the synthesis of organic acid in roots was intensive that of in leaves. The excretion of organic acid and the synthesis of citric acid and malic acid in roots or leaves was a significant positive correlation. These indicated that anion channel and citrate carrier not only regulated excretion of organic acid, but also affected synthesis of organic acid.
The activities of CS, MDH, PEPC and ACO of roots and leaves increased significantly in oilseed rape under single Al stress, the increasing ratios of CS, MDH, PEPC were over 100%, whereas ACO increased less than the three. The trend of change made for accumulation of citric acid and malic acid in roots and leaves. With adding inhibitors into AlCl_3 culture solution, the activities of CS, MDH, PEPC decreased with different degree, ACO was reverse, compared to single Al treatment. These indicated that inhibitor-Al treatments depressed the ability of synthesis of organic acid in plant, and may to citric acid transformed into aconitic acid, thus reduced the contents of citric acid in organization of plant, finally decreased the excretion of organic acid.
The contents of amino acid and soluble sugar increased under Al stress, but it was not significantly difference compared to CK. Complex treatments more increased the osmosis of amino acid and soluble sugar in the root exudation, especially 20μmol/L of A9C that reached markedly level (P<0.05) compared to single Al treatment and CK. The permeability of plasma membrane was increased under Al stress, especially added the A9C and Mersal in the Al solution. It was considered that the osmosis increasing of amino acid and soluble sugar was result of the damage of plasma membrane damage by Al stress. The increasing of pH in excretion under single Al stress was considered an adaptive response on Al stress, and the pH declining after subjoining A9C and Mersal in Al solutions was result in more toxicity of Al to oilseed rape seedlings.
When addition of A9C or Mersal under the Al stress, the dry weights of root and shoot and the ratio of root/shoot decreased rapidly, the growth ratios of main root, plant height and leaf amount inhibited significantly, the root morphology (root length, surface area, average diameter of root, root volume and number of root tips) and moisture contents of root and shoot descended too. In addition, the contents of N, P, K, Ca and Mg in root, stem and leaf of plant declined while Na increased in different extent compared to single Al treatment, which consequently enhanced the ratio of Na~+/ K~+ of root, stem and leaf. Moreover, the accumulation of Al in root, stem and leaf increased obviously with adding A9C and Mersal. It was suggested that A9C and Mersal might decrease the synthesis or excretion of organic acid in oilseed rape, therefore reduced the combine to organic acid with Al, finally the Al toxicity was fierce in plant.
引文
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