植物低温胁迫Ca~（2+）信号的遗传学和细胞学综合分析
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摘要
低温是限制许多植物地理分布和生长的环境因子,严重影响着世界上作物的产量和品质。在拟南芥中鉴定出许多低温信号转导的中间成分,目前对低温信号转导的遗传学研究取得了重要进展。另外,低温刺激之后植物细胞中Ca~(2+)浓度有一个迅速升高的过程,此过程在时间和机制上与低温胁迫感应过程紧密相关。但是,目前对这些基因和低温胁迫中Ca~(2+)信号之间的关系仍不清楚。
为了分析Ca~(2+)信号系统在低温信号转导途径中的作用,我们将水母发光蛋白基因分别转入atfry1、atlos1、atlos5、aticel、atcbl1、athos1、atcipk15、atcbl9-1,等拟南芥突变体和CBF1,CBF2,CBF3超表达拟南芥突变体中,测定冷刺激时细胞中Ca~(2+)浓度的变化。结果显示athos1、atcbl9-1和CBF1超表达材料细胞中Ca~(2+)浓度上升的幅度比对照高,而atfry1、atlos1、atlos5、atice1、atcbl1、atcipk15和CBF2、CBF3超表达等8种材料细胞中Ca~(2+)浓度变化与对照相似。这些结果可以初步帮助我们分析Ca~(2+)信号系统在低温信号转导途径中的位置和作用,并分析这些基因和Ca~(2+) signature的关系。
上述结果表明,冷胁迫刺激之后atcbl9-1中Ca~(2+)浓度变化明显高于野生型,我们进一步分析了atcbl9-1细胞中Ca~(2+)浓度变化和冷胁迫之间的关系。正常生长条件下,atcbl9-1与野生型在表型上没有差别,4℃处理以后二者出现了明显的区别,野生型拟南芥叶片出现严重的玻璃化现象,与atcbl9-1相比,离子渗漏增加,细胞膜的完整性受到损害。对于零下低温atcbl9-1同样表现出较野生型拟南芥更强的抗性,当大部分野生型拟南芥死亡时,仍然有较多的atcbl9-1存活,atcbl9-1的存活率明显高于野生型。这些结果说明atcbl9-1对冷不敏感。
细胞内Ca~(2+)浓度升高可以激活一些转录因子的表达,这些转录因子调控下游许多冷胁迫相关基因的表达,最终提高植物的抗冷性。利用水母发光蛋白检测atcbl9-1中冷刺激时Ca~(2+)浓度变化时发现,冷胁迫刺激条件下atcbl9-1细胞质中Ca~(2+)浓度变化幅度明显高于野生型,而在液泡的胞质面两者Ca~(2+)浓度变化相似。拟南芥预先用Ca~(2+)整合剂EGTA和Ca~(2+)通道抑制剂LaCl_3处理后,atcbl9-1中Ca~(2+)浓度下降幅度大于野生型,利用细胞内钙库的抑制剂LiCl处理后,两者Ca~(2+)浓度下降幅度类似,这些结果暗示,冷胁迫刺激时atcbl9-1内较高的Ca~(2+)浓度可能来源于细胞外的钙库。
AtCBL9序列的N末端包含有一段保守的豆蔻酰化结构域。利用报告基因GFP对AtCBL9进行亚细胞定位,显示AtCBL9蛋白定位于细胞膜上。据此推测AtCBL9可能在质膜上感知外界低温信号,并通过与其相互作用的CIPK蛋白激酶控制下游基因的表达从而调控对低温的应答。为此我们分析了一些受冷诱导基因表达的情况,atcbl9-1中CBF2的表达量比野生型中高,而CBF1和CBF3的表达量却比野生型的低。同时,冷处理条件下atcbl9-1中RD29A,KIN1和COR15A等基因的表达量高于野生型。
总之,定位于质膜上的AtCBL9参与了低温信号转导过程,并且作为COR基因的负调节因子参与了基因的转录调节。
Low temperature is an important environmental factor influencing plant growth and development,and it often extensively affects crop quality and productivity.During the past few years,many intermediates in the cold signal transduction pathway have been identified in Arabidopsis and considerable progress in genetic research has been made in dissecting their functional significance.It has been demonstrated that the earliest reported event in plant responses to low temperature is a transient increase in the concentration of cytosolic free calcium.Despite the fact that many genes are being identified at a rapid pace,the relationships between those genes and the Ca~(2+) are not clear.
To address the relationship between those genes and Ca~(2+),we transformed the atfry1,atlos1, atlos5,atice1,atcbl1,athos1,atcipk15,atcbl9-Imutants and CBF1,CBF2,CBF3 overexpression lines with the plasmid of pMAQ2 and then measured cold-triggered calcium responses.The results demonstrated that cold-triggered calcium responses in athos1,atcbl9-1 mutants and CBF1 overexpression lines were higher than those in control,but were similar in atfry1,atlos1,atlos5, atcbl1,ateipk15,atcbl1,CBF2,CBF3 mutants.These findings could help us define the intermediates of Ca~(2+)-mediated signal transduction into nucleus and their spatial and temporal relationships in cold signaling.
However,at present,little is known about the function of AtCBL9 in cold stress signaling in plants.Here,we report the function of AtCBL9 response to low temperature.Under normal growth conditions,atebl9-1 plants grow like wild-type plants,but chilling injury became apparent after 4℃treatment.The atcbl9-1 leaves showed little increase in electrolyte leakage than the wild type during chilling treatment.The atcbl9-1 mutant plants have shown substantially more tolerant to freezing than wild-type plants under both cold-acclimating and nonacclimating conditions.These results from these studies indicated that atcbl9-1 is more tolerance to cold.
It has been identified that an increase in intracellular calcium can activate the expression of transcription factors that control the subsequent transcription of many genes which must be switched on to provide cold tolerance in the plant.We generated transgenic Arabidopsis plants stably transformed with the plasmids of pMAQ2 and HVA1-2.2.Then,we measured cold-triggered calcium responses in atcbl9-1 and wild-type plants.The atcbl9-1 mutant plants exhibited a higher mean peak[Ca~(2+)]_c than the wild-type plants,but the calcium signature adjacent to the vacuolar membrane rose to similar levels.When Arabidopsis seedlings were pretreated with EGTA or LaCl_3, the mean peak values of wild-type and mutant plants descend to similar levels.LiCl which prevents calcium release from intracellular pools has little effect on the decreased extent.These results may indicate the main involvement of extracellular calcium in this response.
The N-terminal sequence of AtCBL9 harbours a conserved myristoylation motif suggesting a membrane localization of the proteins.The expression pattern at the subcellular level using GFP-AtCBL9 transgenic plants shows the localization of AtCBL9 on the plasma membrane.We speculated that AtCBL9 sensor cold signal in the membrane and regulate the expression of genes whose products are required for stress tolerance.In the atcbl9-1 mutants,transcript levels of CBF2, RD29A,KIN1 and COR15A were substantially higher than in the wild-type.
Taken together,as a Ca~(2+) sensor in the membrane AtCBL9 participates in the cold signal transduction and acts to negatively control the expression of the COR genes.
为了分析Ca~(2+)信号系统在低温信号转导途径中的作用,我们将水母发光蛋白基因分别转入atfry1、atlos1、atlos5、aticel、atcbl1、athos1、atcipk15、atcbl9-1,等拟南芥突变体和CBF1,CBF2,CBF3超表达拟南芥突变体中,测定冷刺激时细胞中Ca~(2+)浓度的变化。结果显示athos1、atcbl9-1和CBF1超表达材料细胞中Ca~(2+)浓度上升的幅度比对照高,而atfry1、atlos1、atlos5、atice1、atcbl1、atcipk15和CBF2、CBF3超表达等8种材料细胞中Ca~(2+)浓度变化与对照相似。这些结果可以初步帮助我们分析Ca~(2+)信号系统在低温信号转导途径中的位置和作用,并分析这些基因和Ca~(2+) signature的关系。
上述结果表明,冷胁迫刺激之后atcbl9-1中Ca~(2+)浓度变化明显高于野生型,我们进一步分析了atcbl9-1细胞中Ca~(2+)浓度变化和冷胁迫之间的关系。正常生长条件下,atcbl9-1与野生型在表型上没有差别,4℃处理以后二者出现了明显的区别,野生型拟南芥叶片出现严重的玻璃化现象,与atcbl9-1相比,离子渗漏增加,细胞膜的完整性受到损害。对于零下低温atcbl9-1同样表现出较野生型拟南芥更强的抗性,当大部分野生型拟南芥死亡时,仍然有较多的atcbl9-1存活,atcbl9-1的存活率明显高于野生型。这些结果说明atcbl9-1对冷不敏感。
细胞内Ca~(2+)浓度升高可以激活一些转录因子的表达,这些转录因子调控下游许多冷胁迫相关基因的表达,最终提高植物的抗冷性。利用水母发光蛋白检测atcbl9-1中冷刺激时Ca~(2+)浓度变化时发现,冷胁迫刺激条件下atcbl9-1细胞质中Ca~(2+)浓度变化幅度明显高于野生型,而在液泡的胞质面两者Ca~(2+)浓度变化相似。拟南芥预先用Ca~(2+)整合剂EGTA和Ca~(2+)通道抑制剂LaCl_3处理后,atcbl9-1中Ca~(2+)浓度下降幅度大于野生型,利用细胞内钙库的抑制剂LiCl处理后,两者Ca~(2+)浓度下降幅度类似,这些结果暗示,冷胁迫刺激时atcbl9-1内较高的Ca~(2+)浓度可能来源于细胞外的钙库。
AtCBL9序列的N末端包含有一段保守的豆蔻酰化结构域。利用报告基因GFP对AtCBL9进行亚细胞定位,显示AtCBL9蛋白定位于细胞膜上。据此推测AtCBL9可能在质膜上感知外界低温信号,并通过与其相互作用的CIPK蛋白激酶控制下游基因的表达从而调控对低温的应答。为此我们分析了一些受冷诱导基因表达的情况,atcbl9-1中CBF2的表达量比野生型中高,而CBF1和CBF3的表达量却比野生型的低。同时,冷处理条件下atcbl9-1中RD29A,KIN1和COR15A等基因的表达量高于野生型。
总之,定位于质膜上的AtCBL9参与了低温信号转导过程,并且作为COR基因的负调节因子参与了基因的转录调节。
Low temperature is an important environmental factor influencing plant growth and development,and it often extensively affects crop quality and productivity.During the past few years,many intermediates in the cold signal transduction pathway have been identified in Arabidopsis and considerable progress in genetic research has been made in dissecting their functional significance.It has been demonstrated that the earliest reported event in plant responses to low temperature is a transient increase in the concentration of cytosolic free calcium.Despite the fact that many genes are being identified at a rapid pace,the relationships between those genes and the Ca~(2+) are not clear.
To address the relationship between those genes and Ca~(2+),we transformed the atfry1,atlos1, atlos5,atice1,atcbl1,athos1,atcipk15,atcbl9-Imutants and CBF1,CBF2,CBF3 overexpression lines with the plasmid of pMAQ2 and then measured cold-triggered calcium responses.The results demonstrated that cold-triggered calcium responses in athos1,atcbl9-1 mutants and CBF1 overexpression lines were higher than those in control,but were similar in atfry1,atlos1,atlos5, atcbl1,ateipk15,atcbl1,CBF2,CBF3 mutants.These findings could help us define the intermediates of Ca~(2+)-mediated signal transduction into nucleus and their spatial and temporal relationships in cold signaling.
However,at present,little is known about the function of AtCBL9 in cold stress signaling in plants.Here,we report the function of AtCBL9 response to low temperature.Under normal growth conditions,atebl9-1 plants grow like wild-type plants,but chilling injury became apparent after 4℃treatment.The atcbl9-1 leaves showed little increase in electrolyte leakage than the wild type during chilling treatment.The atcbl9-1 mutant plants have shown substantially more tolerant to freezing than wild-type plants under both cold-acclimating and nonacclimating conditions.These results from these studies indicated that atcbl9-1 is more tolerance to cold.
It has been identified that an increase in intracellular calcium can activate the expression of transcription factors that control the subsequent transcription of many genes which must be switched on to provide cold tolerance in the plant.We generated transgenic Arabidopsis plants stably transformed with the plasmids of pMAQ2 and HVA1-2.2.Then,we measured cold-triggered calcium responses in atcbl9-1 and wild-type plants.The atcbl9-1 mutant plants exhibited a higher mean peak[Ca~(2+)]_c than the wild-type plants,but the calcium signature adjacent to the vacuolar membrane rose to similar levels.When Arabidopsis seedlings were pretreated with EGTA or LaCl_3, the mean peak values of wild-type and mutant plants descend to similar levels.LiCl which prevents calcium release from intracellular pools has little effect on the decreased extent.These results may indicate the main involvement of extracellular calcium in this response.
The N-terminal sequence of AtCBL9 harbours a conserved myristoylation motif suggesting a membrane localization of the proteins.The expression pattern at the subcellular level using GFP-AtCBL9 transgenic plants shows the localization of AtCBL9 on the plasma membrane.We speculated that AtCBL9 sensor cold signal in the membrane and regulate the expression of genes whose products are required for stress tolerance.In the atcbl9-1 mutants,transcript levels of CBF2, RD29A,KIN1 and COR15A were substantially higher than in the wild-type.
Taken together,as a Ca~(2+) sensor in the membrane AtCBL9 participates in the cold signal transduction and acts to negatively control the expression of the COR genes.
引文
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