盐草（Distichlis spicata）耐盐生理及Na~+转运机理研究
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摘要
本研究着眼于植物耐盐性和Na+转运的研究,对盐生植物——盐草(Distichlis spicata)和甜土植物——高羊茅(Festuca arundinacea)的耐盐生理特性进行了研究,并从个体和细胞水平两个不同的研究层面测定了Na+的分布、转运,分析了各自相关的耐盐机制。研究内容及主要结果概括如下:
1.盐草和高羊茅耐盐生理特性及抗氧化体系反应机制。实验测定了250mM NaCl处理盐草和高羊茅植株0、5、10、15、20天后,其生物量、净光合速率、细胞膜透性、脯氨酸含量以及抗氧化体系的变化。结果表明,随着盐胁迫时间的延长,盐草和高羊茅的生物量和净光合速率都小于对照,但是高羊茅下降的幅度明显大于盐草;盐胁迫导致两种参试材料叶中细胞膜透性增加,高羊茅叶中的细胞膜透性高于盐草;而脯氨酸的含量在两种参试材料中的变化并不明显;盐胁迫导致两种参试材料根组织和叶组织中SOD酶活性增高,而盐草叶组织和根组织中SOD活性高于高羊茅,说明盐草对O2.-有更高的清除效率;盐草叶组织中APX活性显著高于高羊茅,但是根组织中APX活性在两种参试材料之间差异不大,说明APX酶和SOD酶联合变化清除SOD反应产生的H2O2,而且其作用部位主要是植物的叶组织。在盐胁迫条件下,盐草叶组织中CAT酶活性显著升高,而高羊茅叶组织中CAT酶活性变化不大。
2.盐胁迫下Na+在盐草和高羊茅体内不同部位的积累和转运以及相关离子的变化。实验测定了250mM NaCl处理0、5、10、15、20、25天后,盐草和高羊茅根、茎、叶中Na+、K+、Ca2+、Mg2+离子含量变化。盐胁迫会导致盐草和高羊茅根、茎、叶中Na+含量升高,在盐胁迫15天后盐草叶中Na+含量显著低于高羊茅(P<0.05);两种植物叶中K+含量呈下降的趋势;在盐处理25天后,盐草叶中的K+含量高于高羊茅;而高羊茅茎和叶中的Ca2+含量高于盐草,这有利于高羊茅维持这些部位组织正常的代谢功能。通过对K+/Na+离子比的分析发现,盐草根中的K+/Na+离子比小于高羊茅,意味着更多的Na+进入到盐草的根中;而在“根-茎”和“茎-叶”的运输过程中,盐草的K+、Na+运输选择系数高于高羊茅,表明盐草对K+、Na+离子的选择运输具有很强的调控能力。
3.运用X射线电子探针研究了两种耐盐性不同的植物材料盐草和高羊茅在盐胁迫下根、茎、叶中的离子微区分布的状况及其与耐盐适应性的关系。将扫描电镜和X射线能谱仪联用,对盐胁迫20天的盐草和高羊茅不同组织部位进行了点扫描分析,测定了不同部位离子分布状况。盐胁迫下,两种植物根组织中氯离子和钠离子的相对重量都很高,但是高羊茅中柱层中的钠离子和氯离子相对重量都高于盐草;和高羊茅相反,在盐草茎中,大量的钠离子主要分布在韧皮部中,而氯离子则主要存在于木质部中;高羊茅叶组织中钠离子相对重量是盐草叶组织中2倍。说明盐草可以通过根组织对离子的选择性吸收降低进入植物体的有害离子含量,并通过离子在茎中的再次分配降低有害离子进入叶组织,从而避免对叶组织造成较大的离子毒害。显然,矿物离子在植物体内中的微区分布是耐盐机制的一个重要组成部分。
4.利用电子显微技术对盐草叶表面的盐腺结构及分布进行了观察,并研究了盐草泌盐特性的动态变化以及相关的生理生态特性,以建立植物光合作用、蒸腾速率、以及环境因子和盐草泌盐特性之间的关系。对在250mMNaCl溶液中生长了30天的盐草测定了光合速率、蒸腾速率的日变化及泌盐量的日变化,并对所分泌盐分的不同离子含量进行了测定。通过对盐草和大气相对湿度的相关性分析,发现泌盐量(Na、K、Ca、Mg)和大气湿度具有很高的相关性(R>0.843),而盐草在上午9:00时盐分分泌速度最大,达到61.3 umol Na+ g-1f.wth-1。对不同元素的分泌数量排序为:Na>>K>Ca>Mg。
5.利用分子生物学技术获得了盐草液泡膜Na+/H+ antiporter基因中间片段,大约为600bp,通过序列比对发现它和其他植物的相同基因具有很高的同源性,确定为我们希望得到的目的片段。利用X-ray技术分析了盐处理0、5、10、15、20、25天后Na+在盐草细胞不同部位的积累(细胞质、液泡和细胞间质),发现液泡和细胞间质中Na+含量明显增加,表明进入叶片中的Na+主要区隔在液泡中和控制在细胞间质中。同时测定了盐草叶组织质膜和液泡膜H+-ATPase质子泵的活性,发现盐胁迫处理使液泡膜H+-ATPase的活性显著升高,而盐草质膜H+-ATPase活性并没有明显的升高,表明细胞液泡中Na+的积累会激活液泡膜H+-ATPase质子泵的活性。通过Real-time PCR技术检测了盐草液泡膜Na+/H+ antiporter的基因表达随时间的变化,发现随着盐胁迫时间的延长,Na+/H+ antiporter表达量显著升高,说明盐诱导会导致Na+/H+ antiporter超量表达。综合分析发现,随着盐胁迫时间的延长,盐草叶中增加的Na+一部分会积累到细胞液泡中形成区隔化,在区隔化的过程中液泡膜H+-ATPase质子泵的活性增强使离子转运过程的驱动力加大,同时液泡膜Na+/H+ antiporter作为转运载体表达量也会协同增高。细胞间质中Na+也会增高,但是质膜H+-ATPase质子泵并没有明显变化,表明细胞可能通过控制Na+在细胞膜上的通透来抑制Na+进入到细胞质中。
The physiological response of saltgrass(halophyte) and tall fescue (glycophyte) to salt stress, as well as the distribution and transport of Na+, were studied experimentally at both cellar and whole plant levels to better understand the mechanism of salt tolerance and Na+ transport in higher plants. The research work and main results are summarized below.
1. The physiological response and the activity of antioxidative enzymes of saltgrass and tall fescue under salt stress were studied. Plant growth, net photosynthesis, relative electrolyte leakage percentage (RELP), proline content and activity of antioxidative enzymes were measured after 0, 5,10,15,20 days of 250mM NaCl treatment. The results showed that the shoot weight and net photosyntheis of both saltgrass and tall fescue were decreased significantly with prolonged salt stress, with the reduction being more evident in tall fescue than in saltgrass. Salt stress led to an increase in RELP in leaf tissues in both species but it is higher in tall fescue, while no significant change in proline cotent was detected in both species. Salt stress also increased the SOD activity in leaves and roots of both species, with that of saltgrass being higher than in tall fescue, suggesting that the saltgrass had better ability of O2.- radical scavenging. With increased salt stress, leaf APX activity in saltgrass was significantly higher than in tall fescue, but there was no significant difference in root APX between the the two species. These suggested that APX and SOD enzymes jointly contributed to scavenging of H2O2 resulted from SOD reaction and the actitivy took place mainly in leaf tissues. The activity of CAT in leaf tissues of saltgrass increased significantly with increased salt stress, however that of tall fescue was not affected significantly by salt stress.
2. The effects of exogenous NaCl on ion distribution and transport were investigated in both species. Na+, K+,Ca2+,Mg2+ content in root, shoot and leaf were measured after 0, 5,10,15,20 days of 250mM NaCl treatment. The results showed that Na+ content in root, shoot and leaf tissues increased as a response to increased salt stress. After 15 days of NaCl treatment, Na+ content in leaf of saltgrass was significantly higher than that of tall fescue (P<0.05). K+ content decreased in leaves of both species, while that of saltgrass was higher than that Tall fescue after 25 days of NaCl treatment. K+ content in leaf tissue, Ca2+ content in shoot and leaf of tall fescue was higher than in saltgrass, indicating a better condition for tall fescue to maintain the normal metabolic function . K+/Na+ ration in root of saltgrass was lower than that of tall fescue, indicating more Na+ leaked into root tissue.Higher K+-Na+ selectivity was detected in the root-to-shoot and shoot-to-root transport in saltgrass as compared with that in Festuca arundinaces. The results indicated that saltgrass had more control on K-Na selectivity.
3. Microdistribution of mineral ions in roots, stems and leaves of saltgrass and tall fescue was investigated under salt stress using an energy dispersive X-ray microanalyzer (EDX) to explore its relations to salt-tolerance. Plants of the two species were grown hydroponically with 250 mM NaCl . Root tips, stem and leaves were harvested for microanalysis 20 days after transplanting. X-ray percentage of content of atom and the peak of X-ray energy spectra intensity were measured in the sections. It was shown that high Cl and Na X-ray peaks were recorded in root tissues of the both specieswith the X-ray percentages of content of Na and Cl in stelar cells of roots of tall fescue being higher than that of Distichlis spicata. In contrast with tall fescue, saltgrass had relatively high percentage of content Na+ in xylem, and Cl- in phloem. The percentage of Na+ in leaves of tall fescue was 2 times of that of saltgrass.The results suggested that saltgrass regulated the absorption of toxic irons by the mechanism of selective absorption in root, recirculated Na back to root through phloem or retrieval of Na+ from xylem so that less toxic ions were transported to leaf tissues. It was clear that a micro-distribution of mineral ions in plants was involved in the salt tolerance machanism.
4. The micro-structure, distribution of salt glands, salt excretion and the ecophysiological environments in saltgrass were observed using scanning electron microscopyin relation to diurnal changes in net photosynthesis, transpiration rate and other ecophysiological factors. Diurnal dynamics of net photosynthesis, transpiration rate, ionic secretion and ion content were measured after 30 days of NaCl treatment. The results showed that there was generally a positive correlation between the atmosphere humidity and ion secretion during the day (R>0.843), with the excretion rate being the highest at 9:00 AM (61.3μmol Na+ g-1f.wt h-1). The ions could be ranked, according to their excretion rate, as Na>>K>Ca>Mg.
5. A segment of vacuolar Na+/H+ antiporter gene was cloned from saltgrass using PCR and a sequence of 600 bp in length was obtained. The sequence was highly homologous with the Na+/H + antiporter genes from other plants. X-ray microanalysis were used to investigate the Na+ cellar distribution in saltgrass after 0,5,10,15,20,25 days of NaCl treatment. The results showed that Na+ in cytoplasm was compartmented into vacuole and apoplast. Meanwhile the activity of vacuolar H+-ATPase increased after salt treatment, while the activity of plasma H+-ATPase did not increase. And the expression of vacuolar Na+/H+ antiporter increased using real-time PCR. In sum, we found that a part of Na+ which increased in leaf of saltgrass was accumulated in vacuolar and was compartmented. In the proceed on compartment, more energy was supplied for Na+ transport by the increase of vacuolar H+-ATPase, in coordination expression of vacuolar Na+/H+ antiporter as carrier was also increasing. However the activity of H+-ATPase in plasma had not significant change although the Na+ in apoplast increased. It suggested that maybe some ion channels were cotrolled to inhibtated Na+ into cytolplasm.
1.盐草和高羊茅耐盐生理特性及抗氧化体系反应机制。实验测定了250mM NaCl处理盐草和高羊茅植株0、5、10、15、20天后,其生物量、净光合速率、细胞膜透性、脯氨酸含量以及抗氧化体系的变化。结果表明,随着盐胁迫时间的延长,盐草和高羊茅的生物量和净光合速率都小于对照,但是高羊茅下降的幅度明显大于盐草;盐胁迫导致两种参试材料叶中细胞膜透性增加,高羊茅叶中的细胞膜透性高于盐草;而脯氨酸的含量在两种参试材料中的变化并不明显;盐胁迫导致两种参试材料根组织和叶组织中SOD酶活性增高,而盐草叶组织和根组织中SOD活性高于高羊茅,说明盐草对O2.-有更高的清除效率;盐草叶组织中APX活性显著高于高羊茅,但是根组织中APX活性在两种参试材料之间差异不大,说明APX酶和SOD酶联合变化清除SOD反应产生的H2O2,而且其作用部位主要是植物的叶组织。在盐胁迫条件下,盐草叶组织中CAT酶活性显著升高,而高羊茅叶组织中CAT酶活性变化不大。
2.盐胁迫下Na+在盐草和高羊茅体内不同部位的积累和转运以及相关离子的变化。实验测定了250mM NaCl处理0、5、10、15、20、25天后,盐草和高羊茅根、茎、叶中Na+、K+、Ca2+、Mg2+离子含量变化。盐胁迫会导致盐草和高羊茅根、茎、叶中Na+含量升高,在盐胁迫15天后盐草叶中Na+含量显著低于高羊茅(P<0.05);两种植物叶中K+含量呈下降的趋势;在盐处理25天后,盐草叶中的K+含量高于高羊茅;而高羊茅茎和叶中的Ca2+含量高于盐草,这有利于高羊茅维持这些部位组织正常的代谢功能。通过对K+/Na+离子比的分析发现,盐草根中的K+/Na+离子比小于高羊茅,意味着更多的Na+进入到盐草的根中;而在“根-茎”和“茎-叶”的运输过程中,盐草的K+、Na+运输选择系数高于高羊茅,表明盐草对K+、Na+离子的选择运输具有很强的调控能力。
3.运用X射线电子探针研究了两种耐盐性不同的植物材料盐草和高羊茅在盐胁迫下根、茎、叶中的离子微区分布的状况及其与耐盐适应性的关系。将扫描电镜和X射线能谱仪联用,对盐胁迫20天的盐草和高羊茅不同组织部位进行了点扫描分析,测定了不同部位离子分布状况。盐胁迫下,两种植物根组织中氯离子和钠离子的相对重量都很高,但是高羊茅中柱层中的钠离子和氯离子相对重量都高于盐草;和高羊茅相反,在盐草茎中,大量的钠离子主要分布在韧皮部中,而氯离子则主要存在于木质部中;高羊茅叶组织中钠离子相对重量是盐草叶组织中2倍。说明盐草可以通过根组织对离子的选择性吸收降低进入植物体的有害离子含量,并通过离子在茎中的再次分配降低有害离子进入叶组织,从而避免对叶组织造成较大的离子毒害。显然,矿物离子在植物体内中的微区分布是耐盐机制的一个重要组成部分。
4.利用电子显微技术对盐草叶表面的盐腺结构及分布进行了观察,并研究了盐草泌盐特性的动态变化以及相关的生理生态特性,以建立植物光合作用、蒸腾速率、以及环境因子和盐草泌盐特性之间的关系。对在250mMNaCl溶液中生长了30天的盐草测定了光合速率、蒸腾速率的日变化及泌盐量的日变化,并对所分泌盐分的不同离子含量进行了测定。通过对盐草和大气相对湿度的相关性分析,发现泌盐量(Na、K、Ca、Mg)和大气湿度具有很高的相关性(R>0.843),而盐草在上午9:00时盐分分泌速度最大,达到61.3 umol Na+ g-1f.wth-1。对不同元素的分泌数量排序为:Na>>K>Ca>Mg。
5.利用分子生物学技术获得了盐草液泡膜Na+/H+ antiporter基因中间片段,大约为600bp,通过序列比对发现它和其他植物的相同基因具有很高的同源性,确定为我们希望得到的目的片段。利用X-ray技术分析了盐处理0、5、10、15、20、25天后Na+在盐草细胞不同部位的积累(细胞质、液泡和细胞间质),发现液泡和细胞间质中Na+含量明显增加,表明进入叶片中的Na+主要区隔在液泡中和控制在细胞间质中。同时测定了盐草叶组织质膜和液泡膜H+-ATPase质子泵的活性,发现盐胁迫处理使液泡膜H+-ATPase的活性显著升高,而盐草质膜H+-ATPase活性并没有明显的升高,表明细胞液泡中Na+的积累会激活液泡膜H+-ATPase质子泵的活性。通过Real-time PCR技术检测了盐草液泡膜Na+/H+ antiporter的基因表达随时间的变化,发现随着盐胁迫时间的延长,Na+/H+ antiporter表达量显著升高,说明盐诱导会导致Na+/H+ antiporter超量表达。综合分析发现,随着盐胁迫时间的延长,盐草叶中增加的Na+一部分会积累到细胞液泡中形成区隔化,在区隔化的过程中液泡膜H+-ATPase质子泵的活性增强使离子转运过程的驱动力加大,同时液泡膜Na+/H+ antiporter作为转运载体表达量也会协同增高。细胞间质中Na+也会增高,但是质膜H+-ATPase质子泵并没有明显变化,表明细胞可能通过控制Na+在细胞膜上的通透来抑制Na+进入到细胞质中。
The physiological response of saltgrass(halophyte) and tall fescue (glycophyte) to salt stress, as well as the distribution and transport of Na+, were studied experimentally at both cellar and whole plant levels to better understand the mechanism of salt tolerance and Na+ transport in higher plants. The research work and main results are summarized below.
1. The physiological response and the activity of antioxidative enzymes of saltgrass and tall fescue under salt stress were studied. Plant growth, net photosynthesis, relative electrolyte leakage percentage (RELP), proline content and activity of antioxidative enzymes were measured after 0, 5,10,15,20 days of 250mM NaCl treatment. The results showed that the shoot weight and net photosyntheis of both saltgrass and tall fescue were decreased significantly with prolonged salt stress, with the reduction being more evident in tall fescue than in saltgrass. Salt stress led to an increase in RELP in leaf tissues in both species but it is higher in tall fescue, while no significant change in proline cotent was detected in both species. Salt stress also increased the SOD activity in leaves and roots of both species, with that of saltgrass being higher than in tall fescue, suggesting that the saltgrass had better ability of O2.- radical scavenging. With increased salt stress, leaf APX activity in saltgrass was significantly higher than in tall fescue, but there was no significant difference in root APX between the the two species. These suggested that APX and SOD enzymes jointly contributed to scavenging of H2O2 resulted from SOD reaction and the actitivy took place mainly in leaf tissues. The activity of CAT in leaf tissues of saltgrass increased significantly with increased salt stress, however that of tall fescue was not affected significantly by salt stress.
2. The effects of exogenous NaCl on ion distribution and transport were investigated in both species. Na+, K+,Ca2+,Mg2+ content in root, shoot and leaf were measured after 0, 5,10,15,20 days of 250mM NaCl treatment. The results showed that Na+ content in root, shoot and leaf tissues increased as a response to increased salt stress. After 15 days of NaCl treatment, Na+ content in leaf of saltgrass was significantly higher than that of tall fescue (P<0.05). K+ content decreased in leaves of both species, while that of saltgrass was higher than that Tall fescue after 25 days of NaCl treatment. K+ content in leaf tissue, Ca2+ content in shoot and leaf of tall fescue was higher than in saltgrass, indicating a better condition for tall fescue to maintain the normal metabolic function . K+/Na+ ration in root of saltgrass was lower than that of tall fescue, indicating more Na+ leaked into root tissue.Higher K+-Na+ selectivity was detected in the root-to-shoot and shoot-to-root transport in saltgrass as compared with that in Festuca arundinaces. The results indicated that saltgrass had more control on K-Na selectivity.
3. Microdistribution of mineral ions in roots, stems and leaves of saltgrass and tall fescue was investigated under salt stress using an energy dispersive X-ray microanalyzer (EDX) to explore its relations to salt-tolerance. Plants of the two species were grown hydroponically with 250 mM NaCl . Root tips, stem and leaves were harvested for microanalysis 20 days after transplanting. X-ray percentage of content of atom and the peak of X-ray energy spectra intensity were measured in the sections. It was shown that high Cl and Na X-ray peaks were recorded in root tissues of the both specieswith the X-ray percentages of content of Na and Cl in stelar cells of roots of tall fescue being higher than that of Distichlis spicata. In contrast with tall fescue, saltgrass had relatively high percentage of content Na+ in xylem, and Cl- in phloem. The percentage of Na+ in leaves of tall fescue was 2 times of that of saltgrass.The results suggested that saltgrass regulated the absorption of toxic irons by the mechanism of selective absorption in root, recirculated Na back to root through phloem or retrieval of Na+ from xylem so that less toxic ions were transported to leaf tissues. It was clear that a micro-distribution of mineral ions in plants was involved in the salt tolerance machanism.
4. The micro-structure, distribution of salt glands, salt excretion and the ecophysiological environments in saltgrass were observed using scanning electron microscopyin relation to diurnal changes in net photosynthesis, transpiration rate and other ecophysiological factors. Diurnal dynamics of net photosynthesis, transpiration rate, ionic secretion and ion content were measured after 30 days of NaCl treatment. The results showed that there was generally a positive correlation between the atmosphere humidity and ion secretion during the day (R>0.843), with the excretion rate being the highest at 9:00 AM (61.3μmol Na+ g-1f.wt h-1). The ions could be ranked, according to their excretion rate, as Na>>K>Ca>Mg.
5. A segment of vacuolar Na+/H+ antiporter gene was cloned from saltgrass using PCR and a sequence of 600 bp in length was obtained. The sequence was highly homologous with the Na+/H + antiporter genes from other plants. X-ray microanalysis were used to investigate the Na+ cellar distribution in saltgrass after 0,5,10,15,20,25 days of NaCl treatment. The results showed that Na+ in cytoplasm was compartmented into vacuole and apoplast. Meanwhile the activity of vacuolar H+-ATPase increased after salt treatment, while the activity of plasma H+-ATPase did not increase. And the expression of vacuolar Na+/H+ antiporter increased using real-time PCR. In sum, we found that a part of Na+ which increased in leaf of saltgrass was accumulated in vacuolar and was compartmented. In the proceed on compartment, more energy was supplied for Na+ transport by the increase of vacuolar H+-ATPase, in coordination expression of vacuolar Na+/H+ antiporter as carrier was also increasing. However the activity of H+-ATPase in plasma had not significant change although the Na+ in apoplast increased. It suggested that maybe some ion channels were cotrolled to inhibtated Na+ into cytolplasm.
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