PtSOS1、PtAKT1在盐生植物小花碱茅抗盐中的作用
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摘要
小花碱茅(Puccinellia tenuiflora)是一种典型的禾本科碱茅属盐生植物,广泛分布在我国北方地区,是一种重要的优良牧草,具有很强的耐盐碱能力。我们前期研究表明,小花碱茅主要耐盐机制是通过限制根部Na+的单向内流以减少Na+的净吸收,使其维持较高的根系K+、Na+选择性,以促进根系对K+的吸收和转运。但对其根系K+、Na+选择性分子机制仍不清楚。SOS1通过介导根部Na+的外排,以维持植株根系K+吸收和转运,可能小花碱茅SOS1在根系K+、Na+选择性运输中起关键作用；在钾亏缺或盐胁迫条件下,与其它阳离子相比,K+通道蛋白AKT1对于K+具有高度的选择性,并在维持根系K+吸收中起重要作用,或许小花碱茅AKT1在根系K+、Na+选择性吸收中起重要作用。鉴于此,本研究克隆和鉴定了小花碱茅PtSOS1、PtAKT1,并探讨了不同浓度NaCl、KCl以及NaCl与KC1下PtSOS1、PtAKT1的转录丰度和Na+、K+积累的关系,以期揭示PtSOS1、PtAKT1在小花碱茅抗盐中的作用。主要结果如下：
(1) PtSOS1编码1143个氨基酸,C-末端含有一个保守环核苷酸结合域(cyclic nucleotide binding domain),与已知的植物质膜Na+/H+转运蛋白具有较高的同源性(>57%)；PtAKT1编码895个氨基酸,与已知的植物AKT1类K+通道蛋白具有较高的同源性(>60%)。
(2)不同浓度NaCI (25-150mM)下,小花碱茅根中的PtSOS1表达水平和根系根系K+、Na+选择性运输能力(ST值)呈急剧增加趋势。
(3)添加0.1～1mM KCl显著地增加了根中PtSOS1的转录水平和根系ST值,但在5-10mM KCl下根中PtSOS1的转录水平和根系ST值下降。
(4)高盐(150mM NaCl)处理下,随着处理时间的延长,低K+(0.1mM KCl)处理根中PtSOS1表达水平及根系ST值显著低于高K+(5mM KC1)处理。
(5)不同浓度NaCl、KCl以及高盐与低K+或高K+互作下,小花碱茅根中PtSOS1表达水平与根系ST值间呈显著的正相关关系。可见,盐胁迫下,PtSOS1在根系K+、Na+选择性运输中起到关键性的作用。
(6)不同浓度KCl(0～10mM)处理间小花碱茅根中的PtAKT1表达水平与整株K+浓度均无显著差异,这说明在钾亏缺及低钾条件下小花碱茅PtAKT1在维持根系对K+吸收中亦发挥重要作用。
(7)在6-48h,150mM NaCl与0.1或5mM KCl处理的根中PtAKTl表达水平显著高于25mM NaCl与0.1或5mM KCl处理的,但随着处理时间的延长(48～96h),其根中PtAKT1表达水平或整株K+浓度降低,且各处理在48～96h间根中PtAKT1表达水平或整株K+浓度无显著差异。150mM NaCl与0.1或5mMKC1处理6-96h,其根系SA值远高于25mM NaCl与0.1或5mM KCl处理的。进一步研究发现,25mM NaCl与0.1或5mM KC1及150mM NaCl与0.1或5mMKCl处理6-96h,小花碱茅根系K+、Na+选择性吸收能力(SA值)与根中PtAKT1表达水平间呈显著的正相关关系。这说明在盐胁迫下小花碱茅PtAKT1在根系K+、Na+选择性吸收中起关键的作用。
综上所述,PtSOS1编码的质膜Na+/H+逆向转运蛋白在调节小花碱茅根系K+、Na+选择性运输中具有重要的作用,并提出盐胁迫下SOS1通过维持木质部周围薄壁细胞膜的完整性来调节质膜上正常的K和Na转运,从而维持了根系K+、Na+选择性运输的作用模型。此外,在盐胁迫下PtAKT1编码的K+通道蛋白在小花碱茅根系K+、Na+选择性吸收中起关键作用。
P. tenuiflora is a monocotyledonous halophyte found in north China, and used as forage as well as for soil improvement. Our previous studies suggested that restricting unidirectional Na+influx into roots with a strong selectivity for K+over Na+was thought to be contributed to the salt tolerance of P. tenuiflora. However, the molecular mechanisms of selectivity for K+over Na+in the regulation of salt tolerance remains unknown. The plasma membrane Na+/H+antiporter (SOS1) was proved to be a Na+efflux protein and also involved in K+uptake and transport, PtSOS1may be contributed to maintain selective transport capacity for K+over Na+(ST) by roots under salt conditions; plants AKT1-type channels (AKT1) exhibited a high selectivity for K+over other monovalent cations and were reputed to specifically mediate K+uptake by roots under K+deficiency or salt stress, PtAKT1may play an key role in selective absorption capacity for K+over Na+(SA) by roots in P. tenuiflora exposed to salt conditions. In this study, to reveal the role of PtSOS1, PtAKT1in salt tolerance of halophyte P. tenuiflora, we isolated cDNAs for Na+/H+antiporter gene (PtSOS1) and AKT1-type K+channels (PtAKT1) from P. tenuiflora, and Na+and K+accumulations in P. tenuiflora exposed to various NaCl, KCl and NaCl plus different KCl treatments were investigated, respectively. The results are as follows;
(1) PtSOS1consisted of1143amino acid residues with a cyclic nucleotide binding domain located at centrally in the long C-terminal tail and shared higher similarity with the known plasma membrane Na+/H+antiporters(over57%); PtAKT1consists of895amino acid residues and shares high similarity with the known AKT1-type channels (over60%).
(2) The expression levels of PtSOS1in root increased significantly with the increase of external NaCl (25-150mM), accompanied by an increase of selective transport (ST) capacity for K+over Na+by roots.
(3) Transcription levels of PtSOS1in root and ST values increased under0.1-1mM KCl, then declined sharply under5-10mM KCl.
(4) Under150mM NaCl, PtSOS1expression levels in root and ST values at0.1mM KCl was significantly lower than that at5mM KCl with the prolonging of treatment time.
(5) A significantly positive correlation was found between PtSOSI expression levels and ST values in root exposed to150mM NaCl plus0.1or5mM KCl treatments, suggesting that PtSOSI may be the major component of selective transport capacity for K+over Na+under salt conditions.
(6) No significant differences were observed in the expression of PtAKT1in root or whole plant K+concentrations among KCl treatments, indicating that PtAKT1contributes to maintain K+uptake by roots under K+deficiency.
(7) The expression of PtAKTl in root at150mM NaCl plus0.1or5mM KCl was significantly higher than that at25mM NaCl plus0.1or5mM KCl treatment for6-48h.. However, the expression of PtAKTl or whole plant K+concentrations in root was down-regulated by different concentrations of NaCl plus KCl treatments for with the increase of treatment time (48-96h), and there were no significant differences in the expression levels of PtAKT1in root or whole plant K concentrations among treatments for48-96h.. SA values in root at150mM NaCl plus0.1or5mM KCl was significantly higher than that at25mM NaCl plus0.1or5mM KCl treatment for6-96h. Furthermore, a significantly positive correlation was found between selective absorption capacity for K+over Na+values and PtAKT1expression levels in root under various concentrations of NaC1plus KCl treatments, suggesting that PtAKT1may play an important role in selective absorption capacity for K+over+Na by roots in P. tenuiflora exposed to salt conditions.
In summary, we proposed that PtSOS1seems likely to be the major component of selective transport capacity for K+over Na+and hence salt tolerance of P. tenuiflora. Finally, we hypothesized a function model of SOS1in regulating K+and Na+transport system in the membrane of xylem parenchyma cells by sustaining the membrane integrity under salt stress. In addition, PtAKT1may contribute to maintaining selective absorption capacity for K+over Na+by roots in response to salt stress.
(1) PtSOS1编码1143个氨基酸,C-末端含有一个保守环核苷酸结合域(cyclic nucleotide binding domain),与已知的植物质膜Na+/H+转运蛋白具有较高的同源性(>57%)；PtAKT1编码895个氨基酸,与已知的植物AKT1类K+通道蛋白具有较高的同源性(>60%)。
(2)不同浓度NaCI (25-150mM)下,小花碱茅根中的PtSOS1表达水平和根系根系K+、Na+选择性运输能力(ST值)呈急剧增加趋势。
(3)添加0.1～1mM KCl显著地增加了根中PtSOS1的转录水平和根系ST值,但在5-10mM KCl下根中PtSOS1的转录水平和根系ST值下降。
(4)高盐(150mM NaCl)处理下,随着处理时间的延长,低K+(0.1mM KCl)处理根中PtSOS1表达水平及根系ST值显著低于高K+(5mM KC1)处理。
(5)不同浓度NaCl、KCl以及高盐与低K+或高K+互作下,小花碱茅根中PtSOS1表达水平与根系ST值间呈显著的正相关关系。可见,盐胁迫下,PtSOS1在根系K+、Na+选择性运输中起到关键性的作用。
(6)不同浓度KCl(0～10mM)处理间小花碱茅根中的PtAKT1表达水平与整株K+浓度均无显著差异,这说明在钾亏缺及低钾条件下小花碱茅PtAKT1在维持根系对K+吸收中亦发挥重要作用。
(7)在6-48h,150mM NaCl与0.1或5mM KCl处理的根中PtAKTl表达水平显著高于25mM NaCl与0.1或5mM KCl处理的,但随着处理时间的延长(48～96h),其根中PtAKT1表达水平或整株K+浓度降低,且各处理在48～96h间根中PtAKT1表达水平或整株K+浓度无显著差异。150mM NaCl与0.1或5mMKC1处理6-96h,其根系SA值远高于25mM NaCl与0.1或5mM KCl处理的。进一步研究发现,25mM NaCl与0.1或5mM KC1及150mM NaCl与0.1或5mMKCl处理6-96h,小花碱茅根系K+、Na+选择性吸收能力(SA值)与根中PtAKT1表达水平间呈显著的正相关关系。这说明在盐胁迫下小花碱茅PtAKT1在根系K+、Na+选择性吸收中起关键的作用。
综上所述,PtSOS1编码的质膜Na+/H+逆向转运蛋白在调节小花碱茅根系K+、Na+选择性运输中具有重要的作用,并提出盐胁迫下SOS1通过维持木质部周围薄壁细胞膜的完整性来调节质膜上正常的K和Na转运,从而维持了根系K+、Na+选择性运输的作用模型。此外,在盐胁迫下PtAKT1编码的K+通道蛋白在小花碱茅根系K+、Na+选择性吸收中起关键作用。
P. tenuiflora is a monocotyledonous halophyte found in north China, and used as forage as well as for soil improvement. Our previous studies suggested that restricting unidirectional Na+influx into roots with a strong selectivity for K+over Na+was thought to be contributed to the salt tolerance of P. tenuiflora. However, the molecular mechanisms of selectivity for K+over Na+in the regulation of salt tolerance remains unknown. The plasma membrane Na+/H+antiporter (SOS1) was proved to be a Na+efflux protein and also involved in K+uptake and transport, PtSOS1may be contributed to maintain selective transport capacity for K+over Na+(ST) by roots under salt conditions; plants AKT1-type channels (AKT1) exhibited a high selectivity for K+over other monovalent cations and were reputed to specifically mediate K+uptake by roots under K+deficiency or salt stress, PtAKT1may play an key role in selective absorption capacity for K+over Na+(SA) by roots in P. tenuiflora exposed to salt conditions. In this study, to reveal the role of PtSOS1, PtAKT1in salt tolerance of halophyte P. tenuiflora, we isolated cDNAs for Na+/H+antiporter gene (PtSOS1) and AKT1-type K+channels (PtAKT1) from P. tenuiflora, and Na+and K+accumulations in P. tenuiflora exposed to various NaCl, KCl and NaCl plus different KCl treatments were investigated, respectively. The results are as follows;
(1) PtSOS1consisted of1143amino acid residues with a cyclic nucleotide binding domain located at centrally in the long C-terminal tail and shared higher similarity with the known plasma membrane Na+/H+antiporters(over57%); PtAKT1consists of895amino acid residues and shares high similarity with the known AKT1-type channels (over60%).
(2) The expression levels of PtSOS1in root increased significantly with the increase of external NaCl (25-150mM), accompanied by an increase of selective transport (ST) capacity for K+over Na+by roots.
(3) Transcription levels of PtSOS1in root and ST values increased under0.1-1mM KCl, then declined sharply under5-10mM KCl.
(4) Under150mM NaCl, PtSOS1expression levels in root and ST values at0.1mM KCl was significantly lower than that at5mM KCl with the prolonging of treatment time.
(5) A significantly positive correlation was found between PtSOSI expression levels and ST values in root exposed to150mM NaCl plus0.1or5mM KCl treatments, suggesting that PtSOSI may be the major component of selective transport capacity for K+over Na+under salt conditions.
(6) No significant differences were observed in the expression of PtAKT1in root or whole plant K+concentrations among KCl treatments, indicating that PtAKT1contributes to maintain K+uptake by roots under K+deficiency.
(7) The expression of PtAKTl in root at150mM NaCl plus0.1or5mM KCl was significantly higher than that at25mM NaCl plus0.1or5mM KCl treatment for6-48h.. However, the expression of PtAKTl or whole plant K+concentrations in root was down-regulated by different concentrations of NaCl plus KCl treatments for with the increase of treatment time (48-96h), and there were no significant differences in the expression levels of PtAKT1in root or whole plant K concentrations among treatments for48-96h.. SA values in root at150mM NaCl plus0.1or5mM KCl was significantly higher than that at25mM NaCl plus0.1or5mM KCl treatment for6-96h. Furthermore, a significantly positive correlation was found between selective absorption capacity for K+over Na+values and PtAKT1expression levels in root under various concentrations of NaC1plus KCl treatments, suggesting that PtAKT1may play an important role in selective absorption capacity for K+over+Na by roots in P. tenuiflora exposed to salt conditions.
In summary, we proposed that PtSOS1seems likely to be the major component of selective transport capacity for K+over Na+and hence salt tolerance of P. tenuiflora. Finally, we hypothesized a function model of SOS1in regulating K+and Na+transport system in the membrane of xylem parenchyma cells by sustaining the membrane integrity under salt stress. In addition, PtAKT1may contribute to maintaining selective absorption capacity for K+over Na+by roots in response to salt stress.
引文
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