碱地肤抗盐碱胁迫的生理机制研究
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摘要
土壤盐碱化已成为一个世界性环境问题,土壤盐化与碱化往往相伴发生。盐胁迫和碱胁迫实际是两种不同性质的胁迫。碱胁迫比盐胁迫更复杂,具有更大的生态破坏力。本文以抗碱盐生植物碱地肤为材料,系统地探讨其抗盐碱生境的生理机制,尤其探讨了草酸积累、分布及其对碱地肤适应盐碱条件的生理贡献,以此为治理盐碱化生境提供重要的理论依据。
一、盐碱胁迫下碱地肤体内渗透调节和离子平衡机制
本文对碱地肤幼苗进行非胁迫和盐胁迫(盐度:200 mM,NaCl: Na_2SO_4 =1:1)或碱胁迫(盐度:200 mM,NaHCO_3: Na_2CO_3 = 1:1)处理。通过观测盐、碱胁迫后溶质积累变化及其分布差异,探讨了盐、碱胁迫下碱地肤的渗透调节和离子平衡机制的动态进程及其分布差异,更加深入地揭示了碱地肤响应盐碱胁迫的生理机制特点。碱地肤体内各种溶质对渗透调节或离子平衡的贡献各不相同,而且在盐碱胁迫下其贡献也发生动态变化。但无论如何,Na~+、K~+、有机酸、Cl-是盐胁迫下主要的渗透调节剂,而Na~+、K~+、有机酸是碱胁迫下主要的渗透调节剂。无论盐、碱胁迫,Na~+、K~+均是正电荷的主要贡献者,而有机酸为负电荷主要提供者。
碱地肤的渗透调节、离子平衡同样存在部位差异。但共同之处是:K~+、Na~+和有机酸是碱地肤各部位主要的渗透调节物质。碱地肤不同部位正电荷主要来源于Na~+、K~+,负电荷主要来源于有机酸。值得注意的是,有机酸对渗透调节和离子平衡的贡献均主要集中体现在叶片。有机酸优先在决定光合生产力的叶片中发挥其生理作用,可以保证整个植株正常的生理代谢。此外,有机酸在盐、碱胁迫下的作用也存在差异,盐胁迫下有机酸作用较弱。这是由于碱地肤此时选择了耗能更少的Cl-和SO42-等无机离子进行离子平衡和渗透调节,可以认为无机离子选择性的吸收是碱地肤适应盐胁迫的重要生存策略,而有机酸的积累是其响应碱胁迫的关键性生理机制之一。
二、盐碱胁迫下碱地肤体内有机酸代谢机制
通过分析碱地肤体内有机酸组分的动态积累及其体内的分布差异,以探讨盐碱胁迫下碱地肤体内有机酸代谢调节机制特点。结果表明,在响应碱胁迫时,碱地肤不同部位也存在着不同的有机酸代谢调节方式。草酸主要分布在茎叶尤其是成熟叶片中;苹果酸为老茎中最主要有机酸;而琥珀酸为根中的主导有机酸。尽管如此,从整个植株来看,草酸仍然是其体内有机酸的主要成分,同时也是盐、碱胁迫下积累的主要有机酸。实验结果还表明,碱地肤根系不分泌草酸。综上可知,草酸只在碱地肤体内特别是决定光合生产力的成熟叶才发挥其生理作用,以保障植株的正常生理代谢。草酸为主有机酸的积累是决定碱地肤既抗盐又抗碱这一适应特性的内在因素之一。
三、盐碱胁迫下草酸的生理贡献及其代谢特点
通过分析草酸的生理作用及其相关代谢酶的活性变化,进一步明确草酸对其适应盐碱生境的生理贡献及其代谢特点。结果表明:草酸以小分子有机阴离子的形式存在于碱地肤体内,不仅对渗透调节和pH调节具有决定作用,也是在维持离子平衡中起主导作用的负电荷供体。实验还表明,碱地肤体内草酸的含量与其分解作用(OXO途径)并不大,而是主要取决于其合成作用。在外施盐、碱胁迫24h后,乙醇酸氧化酶(GO: EC1.1.3.1),异柠檬酸裂解酶(ICL: EC 4.1.3.1)和PEP羧化酶(PEPcase: EC 4.1.1.31)的活性均显著升高,通过相关分析的结果表明,GO才是合成草酸的关键酶。因此证明了GO才是决定草酸积累的关键限制因子。
碱地肤在进化中形成了一套适应盐、碱胁迫的生理机制,即主要通过提高GO关键酶的活性来调动草酸为主的有机酸的代谢调节,以达到渗透调节、离子平衡及调节pH值的综合目的。
Soil salinization has become a global environmental problem. Soil alkalization and salinization frequently co-occur. The alkali stress and salt stress are two distinct kinds of stresses actually. Alkali-stress more complex and serious destructive effect on ecological environment than that salt-stress. In present study, we chosen an alkali-tolerant halophyte Kochia sieversiana as the test organism, probed the physiological mechanisms of K. sieversiana to salt and alkali stresses, especially, we revealed oxalic acid accumulation and distribution of K. sieversiana and its physiological contribution to the adaptability of K sieversiana to saline and alkaline conditions. On the basis of these improved the theoretical basis for the treatment of salt-alkalinized habitats.
1. The mechanisms of ion balance and osmotic adjustment in Kochia sieversiana under salt- and alkali-stress
In the present study, seedlings of Kochia sieversiana were exposed to the following conditions: non-stress, salt stress (molar ratio of NaCl: Na_2SO_4= 1:1, salinity: 200mM) and alkali stress (molar ratio of NaHCO_3: Na_2CO_3= 1:1, salinity: 200mM). The solutes accumulation and the differences in solutes distribution under salt- or alkali- stress were monitored in order to investigate the dynamic process and distribution differences of osmotic adjustment and ion balance in K. sieversiana during responding to salt and alkali stresses. Our work revealed the physiological mechanisms in K. sieversiana responding to salt and alkali stresses more deeply.
The contributions of different solutes to osmotic adjustment and ion balance were different, and their contributions under salt- or alkali- stress were discrepant too. But anyhow Na~+, K~+, organic acid, and Cl~- were the main osmolytes under salt stress, while Na~+, K~+, organic acid were the main osmolytes under alkali stress. Whether under salt stress or under alkali stresses, Na~+ and K~+ were the main contributors to positive charge, and organic acid was the dominant contributor to negative charge.
There were differences among different parts of K. sieversiana in the osmotic adjustment and ion balance, but the common point: K~+, Na~+ and organic acid were the main osmolytes in different parts of K. sieversiana. The positive charge mainly came from Na~+ and K~+, and the negative charge mainly came from organic acid in different parts. Remarkably, the contributions of organic acid to osmotic adjustment and ion balance was concentrated in leaves. Organic acid play physiological role preferentially in leaves, on which photosynthesis productivity was based, and ensured to maintain normal physiological metabolism in whole plant. The effects of organic acid on salt and alkali stresses were different, and its effect was weaker under salt stress. That was because that Cl- or SO42- were accumulated selectively in shoots, which accumulation process consumed less energy. The selective absorption of inorganic ions in K. sieversiana to adapt to salt stress is an important survival strategy, while the accumulation of organic acids was one of the key
2. The mechanisms of organic acids metabolism in K. sieversiana under salt and alkali stresses
The characteristic of mechanisms of organic acids metabolism under salt and alkali stresses were discussed by analyzing the dynamic accumulation and differences of distribution of organic acids in K. sieversiana.
The results showed that different parts of K. sieversiana differed in organic acids metabolic regulation mechanisms of adaptability to alkali conditions. Oxalic acid was the main constituent in shoots, especially mature leaves; malic acid was the main organic acid in old stems; and succinic acid was the dominant organic acid in roots. However, for the whole plant, oxalic acid was the main organic acid observed in plants, and it was also the main organic acid observed in plants under either salt or alkali stress. The root secretion experiment indicated that oxalic acid was not detected in root exudates. Oxalic acid played its physiological action in functional leaves on which photosynthesis productivity is dependent on, and ensured plant had normal physiological metabolic. The accumulation of oxalic acid was one of the significant endogenous factors which determined the adaptive characteristics of both salt-resistance and alkali-resistance for K. sieversiana.
3. Physiological contribution of oxalic acid and the characteristics of its metabolism in K. sieversiana under salt and alkali stresses
The contribution of oxalic acid to salt-alkalinized soil adaptability and the characteristic its metabolism were identified by studying the physiological action of oxalic acid and change of its related enzyme activities. The results showed that oxalic acid anion was not only an organic osmotic regulator, but also a negative charge donator, playing leading roles in maintaining ion balance and pH adjustment in vivo. Results also showed that the oxalic acid content in K. sieversiana and its decomposition (OXO way) were not entirely relevant, but mainly depended on its synthesis. K. sieversiana were exposed to salt stress and alkali stress after 24 hours, GO, ICL and PEPcase activities increased significantly. The analysis of correlation showed that GO was a key enzyme for oxalic acid biosynthesis. Therefore, this proved that GO was the key limiting factor for the accumulation of oxalic acid.
K. sieversiana in evolution formed physiological mechanisms to adaption of alkali and salt stresses, mainly by increasing the activity of key enzyme GO to mobilize the metabolism of oxalic acid-based regulation, in order to achieve comprehensive purposes of osmotic adjustment, ion balance and pH regulation.
一、盐碱胁迫下碱地肤体内渗透调节和离子平衡机制
本文对碱地肤幼苗进行非胁迫和盐胁迫(盐度:200 mM,NaCl: Na_2SO_4 =1:1)或碱胁迫(盐度:200 mM,NaHCO_3: Na_2CO_3 = 1:1)处理。通过观测盐、碱胁迫后溶质积累变化及其分布差异,探讨了盐、碱胁迫下碱地肤的渗透调节和离子平衡机制的动态进程及其分布差异,更加深入地揭示了碱地肤响应盐碱胁迫的生理机制特点。碱地肤体内各种溶质对渗透调节或离子平衡的贡献各不相同,而且在盐碱胁迫下其贡献也发生动态变化。但无论如何,Na~+、K~+、有机酸、Cl-是盐胁迫下主要的渗透调节剂,而Na~+、K~+、有机酸是碱胁迫下主要的渗透调节剂。无论盐、碱胁迫,Na~+、K~+均是正电荷的主要贡献者,而有机酸为负电荷主要提供者。
碱地肤的渗透调节、离子平衡同样存在部位差异。但共同之处是:K~+、Na~+和有机酸是碱地肤各部位主要的渗透调节物质。碱地肤不同部位正电荷主要来源于Na~+、K~+,负电荷主要来源于有机酸。值得注意的是,有机酸对渗透调节和离子平衡的贡献均主要集中体现在叶片。有机酸优先在决定光合生产力的叶片中发挥其生理作用,可以保证整个植株正常的生理代谢。此外,有机酸在盐、碱胁迫下的作用也存在差异,盐胁迫下有机酸作用较弱。这是由于碱地肤此时选择了耗能更少的Cl-和SO42-等无机离子进行离子平衡和渗透调节,可以认为无机离子选择性的吸收是碱地肤适应盐胁迫的重要生存策略,而有机酸的积累是其响应碱胁迫的关键性生理机制之一。
二、盐碱胁迫下碱地肤体内有机酸代谢机制
通过分析碱地肤体内有机酸组分的动态积累及其体内的分布差异,以探讨盐碱胁迫下碱地肤体内有机酸代谢调节机制特点。结果表明,在响应碱胁迫时,碱地肤不同部位也存在着不同的有机酸代谢调节方式。草酸主要分布在茎叶尤其是成熟叶片中;苹果酸为老茎中最主要有机酸;而琥珀酸为根中的主导有机酸。尽管如此,从整个植株来看,草酸仍然是其体内有机酸的主要成分,同时也是盐、碱胁迫下积累的主要有机酸。实验结果还表明,碱地肤根系不分泌草酸。综上可知,草酸只在碱地肤体内特别是决定光合生产力的成熟叶才发挥其生理作用,以保障植株的正常生理代谢。草酸为主有机酸的积累是决定碱地肤既抗盐又抗碱这一适应特性的内在因素之一。
三、盐碱胁迫下草酸的生理贡献及其代谢特点
通过分析草酸的生理作用及其相关代谢酶的活性变化,进一步明确草酸对其适应盐碱生境的生理贡献及其代谢特点。结果表明:草酸以小分子有机阴离子的形式存在于碱地肤体内,不仅对渗透调节和pH调节具有决定作用,也是在维持离子平衡中起主导作用的负电荷供体。实验还表明,碱地肤体内草酸的含量与其分解作用(OXO途径)并不大,而是主要取决于其合成作用。在外施盐、碱胁迫24h后,乙醇酸氧化酶(GO: EC1.1.3.1),异柠檬酸裂解酶(ICL: EC 4.1.3.1)和PEP羧化酶(PEPcase: EC 4.1.1.31)的活性均显著升高,通过相关分析的结果表明,GO才是合成草酸的关键酶。因此证明了GO才是决定草酸积累的关键限制因子。
碱地肤在进化中形成了一套适应盐、碱胁迫的生理机制,即主要通过提高GO关键酶的活性来调动草酸为主的有机酸的代谢调节,以达到渗透调节、离子平衡及调节pH值的综合目的。
Soil salinization has become a global environmental problem. Soil alkalization and salinization frequently co-occur. The alkali stress and salt stress are two distinct kinds of stresses actually. Alkali-stress more complex and serious destructive effect on ecological environment than that salt-stress. In present study, we chosen an alkali-tolerant halophyte Kochia sieversiana as the test organism, probed the physiological mechanisms of K. sieversiana to salt and alkali stresses, especially, we revealed oxalic acid accumulation and distribution of K. sieversiana and its physiological contribution to the adaptability of K sieversiana to saline and alkaline conditions. On the basis of these improved the theoretical basis for the treatment of salt-alkalinized habitats.
1. The mechanisms of ion balance and osmotic adjustment in Kochia sieversiana under salt- and alkali-stress
In the present study, seedlings of Kochia sieversiana were exposed to the following conditions: non-stress, salt stress (molar ratio of NaCl: Na_2SO_4= 1:1, salinity: 200mM) and alkali stress (molar ratio of NaHCO_3: Na_2CO_3= 1:1, salinity: 200mM). The solutes accumulation and the differences in solutes distribution under salt- or alkali- stress were monitored in order to investigate the dynamic process and distribution differences of osmotic adjustment and ion balance in K. sieversiana during responding to salt and alkali stresses. Our work revealed the physiological mechanisms in K. sieversiana responding to salt and alkali stresses more deeply.
The contributions of different solutes to osmotic adjustment and ion balance were different, and their contributions under salt- or alkali- stress were discrepant too. But anyhow Na~+, K~+, organic acid, and Cl~- were the main osmolytes under salt stress, while Na~+, K~+, organic acid were the main osmolytes under alkali stress. Whether under salt stress or under alkali stresses, Na~+ and K~+ were the main contributors to positive charge, and organic acid was the dominant contributor to negative charge.
There were differences among different parts of K. sieversiana in the osmotic adjustment and ion balance, but the common point: K~+, Na~+ and organic acid were the main osmolytes in different parts of K. sieversiana. The positive charge mainly came from Na~+ and K~+, and the negative charge mainly came from organic acid in different parts. Remarkably, the contributions of organic acid to osmotic adjustment and ion balance was concentrated in leaves. Organic acid play physiological role preferentially in leaves, on which photosynthesis productivity was based, and ensured to maintain normal physiological metabolism in whole plant. The effects of organic acid on salt and alkali stresses were different, and its effect was weaker under salt stress. That was because that Cl- or SO42- were accumulated selectively in shoots, which accumulation process consumed less energy. The selective absorption of inorganic ions in K. sieversiana to adapt to salt stress is an important survival strategy, while the accumulation of organic acids was one of the key
2. The mechanisms of organic acids metabolism in K. sieversiana under salt and alkali stresses
The characteristic of mechanisms of organic acids metabolism under salt and alkali stresses were discussed by analyzing the dynamic accumulation and differences of distribution of organic acids in K. sieversiana.
The results showed that different parts of K. sieversiana differed in organic acids metabolic regulation mechanisms of adaptability to alkali conditions. Oxalic acid was the main constituent in shoots, especially mature leaves; malic acid was the main organic acid in old stems; and succinic acid was the dominant organic acid in roots. However, for the whole plant, oxalic acid was the main organic acid observed in plants, and it was also the main organic acid observed in plants under either salt or alkali stress. The root secretion experiment indicated that oxalic acid was not detected in root exudates. Oxalic acid played its physiological action in functional leaves on which photosynthesis productivity is dependent on, and ensured plant had normal physiological metabolic. The accumulation of oxalic acid was one of the significant endogenous factors which determined the adaptive characteristics of both salt-resistance and alkali-resistance for K. sieversiana.
3. Physiological contribution of oxalic acid and the characteristics of its metabolism in K. sieversiana under salt and alkali stresses
The contribution of oxalic acid to salt-alkalinized soil adaptability and the characteristic its metabolism were identified by studying the physiological action of oxalic acid and change of its related enzyme activities. The results showed that oxalic acid anion was not only an organic osmotic regulator, but also a negative charge donator, playing leading roles in maintaining ion balance and pH adjustment in vivo. Results also showed that the oxalic acid content in K. sieversiana and its decomposition (OXO way) were not entirely relevant, but mainly depended on its synthesis. K. sieversiana were exposed to salt stress and alkali stress after 24 hours, GO, ICL and PEPcase activities increased significantly. The analysis of correlation showed that GO was a key enzyme for oxalic acid biosynthesis. Therefore, this proved that GO was the key limiting factor for the accumulation of oxalic acid.
K. sieversiana in evolution formed physiological mechanisms to adaption of alkali and salt stresses, mainly by increasing the activity of key enzyme GO to mobilize the metabolism of oxalic acid-based regulation, in order to achieve comprehensive purposes of osmotic adjustment, ion balance and pH regulation.
引文
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