小麦CBL基因和CIPK基因的克隆及在非生物胁迫中的功能研究
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摘要
小麦是世界上最重要的粮食作物,其种植面积、总产量及总贸易额均居粮食作物之首。干旱、高盐等是严重影响小麦产量和品质的非生物逆境,研究小麦对逆境信号的感知、传导以及抗逆性的分子机制,对小麦抗逆分子育种研究与开发具有重要意义。Ca2+作为第二信使介导植物对各种不同逆境胁迫的反应,这其中涉及到复杂的信号转导途径。类钙调磷酸酶B蛋白(Calcineurin B-Like, CBL)和其互作蛋白激酶(CBL-interacting protein kinase, CIPK)是与非生物逆境胁迫相关的Ca2+信号转导中很重要的组分。然而到目前为止,除了发现一个小麦CIPK基因(WPK4)参与细胞分裂素以及光和营养胁迫信号转导外,对小麦CBL和CIPK基因的系统研究还未见报道。我们在对DFCI数据库和NCBI数据库中小麦CBL和CIPK基因相关EST序列的搜索和分析的基础上,利用RACE技术分别成功地克隆了7个CBL基因和8个CIPK基因,并对它们编码的蛋白质进行了系统的分析。利用酵母双杂交方法研究了CBL和CIPK之间特异的相互作用。利用RT-PCR和qRT-PCR技术分析了这些基因在不同非生物胁迫逆境处理下的表达水平。另外,进一步通过对转TaCIPK14基因和转TaCIPK29基因的烟草在非生物胁迫下的表型,生理生化指标与分子机制的分析,揭示了TaCIPK14和TaCIPK29在非生物胁迫中功能和作用机理,取得的主要研究结果如下：
1)小麦CBL和CIPK蛋白与水稻同源的CBL和CIPK蛋白的氨基酸序列和结构高度相似,这些CBL蛋白均含有四个能结合Ca2+的EF-hands结构域和一个FPSF结构域；CIPK蛋白都含有激酶结构域,FISL结构域和PPI结构域。进化分析显示,小麦CBL蛋白和CIPK蛋白和水稻同源CBL蛋白和CIPK蛋白都归类于相同的亚族。
2)基因表达分析结果表明,小麦特定的CBL或CIPK基因对冷、盐、渗透胁迫和ABA处理的响应模式不同,而且同一处理能够被不同的CBL或CIPK基因感应,这说明小麦CBL和CIPK基因可能介导对不同胁迫信号的交联过程。
3)利用酵母双杂交技术分析发现,小麦不同的CBL和CIPK蛋白之间的相互作用的特异性和强度不同。一些重要的特异性的CBL-CIPK互作子如：TaCBL1-TaCIPK9, TaCBL1-TaCIPK23, TaCBL2-TaCIPK2, TaCBL3-TaCIPK2, TaCBL2-TaCIPK29和TaCBL3-TaCIPK29可能参与小麦对不同非生物胁迫的响应。这些结果表明,小麦CBL和CIPK对不同非生物胁迫的信号转导是通过形成不同的CBL-CIPK复合物来解码特定胁迫下产生的特异Ca2+信号。
4)系统分析了TaCIPK14基因和TaCIPK29基因在非生物胁迫反应中的作用。TaCIPK14基因受冷、渗透胁迫和盐胁迫等非生物胁迫以及胁迫相关信号分子ABA、乙烯和H202诱导。证明了在烟草中过表达TaCIPK14基因可提高转基因烟草植株对冷和盐胁迫的抗性。TaCIPK14提高对冷和盐的抗性是通过调控一些抗氧化酶或胁迫相关基因如NtCAT, NtDREB3和NtLEA等的表达来实现的。增强的抗氧化酶的活性有助于清除逆境下产生的过量ROS,减轻ROS对细胞膜的损伤。同时,TaCIPK14能降低转基因烟草在盐胁迫下的Na+含量和提高细胞的K+/Na+比率,这也有助于植物对盐胁迫的抗性。
5) TaCIPK29基因能够被盐、冷、甲基紫晶(MV)、ABA和乙烯诱导。过表达TaCIPK29基因的烟草植株增强了对盐胁迫的抗性。TaCIPK29通过提高一些通道蛋白基因如NtSOS1, NtNHX2, NtNHX4和NtCAX3的表达来维持高的K+/Na+比和Ca2+含量以及通过增强CAT和POD的表达和活性来减少H2O2对细胞膜的损伤来抵抗盐胁迫的伤害。TaCIPK29没有特定的定位信号,在整个细胞中都有分布。。TaCIPK29不仅能够特异地与小麦的TaCBL2和TaCBL3互作,也能与烟草的NtCBL2和NtCBL3互作,说明TaCIPK29通过CBL2和CBL3传递Ca2+信号。同时TaCIPK29还能与烟草的NtCAT1相互作用,增强其活性,这有助于清除过多的H202。因此,TaCIPK29是一个盐胁迫中的正调控因子,参与了盐胁迫下对离子和ROS的调控。
综上所述,如同水稻和拟南芥中同源的CBL和CIPK基因一样,小麦CBL和CIPK基因也是非生物胁迫响应基因。不同的CBL和CIPK基因可能赋予植物对不同逆境的抗性。已鉴定的抗逆基因TaCIPK14和TaCIPK29可作为培育转基因抗逆小麦新品种的候选基因。本研究为深入研究小麦中这两个重要的基因家族的功能和分子机制奠定了基础,同时也有助于小麦抗逆分子育种研究与开发。
Wheat is a world staple crop, and its acreage, total production and trade ranks the first place among food crops. However, wheat production is often constrained by various abiotic stresses, such as drought, salinity, and extreme temperatures. Understanding the molecular mechanism of the abiotic stress responses can facilitate the genetic improvement of stress tolerance in wheat. Calcium, as a second messenger, is involved in the mediation of various responses to different environmental stresses and related to highly complex signal transduction pathways. Calcineurin B-like proteins (CBLs) and their target proteins, the CBL-interacting protein kinases (CIPKs), have emerged as key Ca2+-mediated signaling components in response to various abiotic stresses in many plants. However, the CBL and CIPK genes in wheat have not yet been comprehensively studied to date, except one CIPK gene (WPK4) which has been reported to mediate cytokine signaling transduction and the response to light and nutrient deprivation. In this study, seven CBL genes and eight CIPK genes were amplified from wheat genome using the RACE technique and their preferential interaction and differential responses to various abiotic stresses were investigated. The roles of two wheat CIPK genes TaCIPK14and TaCIPK29in response to abiotic stresses were further investigated. The main results are as follows:
1) Wheat CBLs and CIPKs were found to be similar to their counterparts in rice in motif structure and subgroup classification. These CBL proteins have four EF-hands and FPSF domains, while CIPK proteins contain kinase domain, FISL domain and PPI domain.
2) The isolated wheat CBL and CIPK genes were found to be expressed differentially in various tissues and in response to different abiotic stresses including cold, salt, and osmotic stresses and exposure to the phytohormone abscisic acid (ABA). Furthermore, we also found that one CBL or CIPK gene was able to respond to several treatments, and one treatment was sensed by multiple CBL or CIPK genes. Thus, TaCBLs and TaCIPKs were found to mediate crosstalk among different signaling pathways.
3) The preferential interactions of TaCBLs and TaCIPKs were identified using a yeast two-hybrid assay. Several important, specific CBL-CIPK interaction partners (TaCBL1-TaCIPK9, TaCBL1-TaCIPK23, TaCBL2-TaCIPK2, and TaCBL3-TaCIPK2, TaCBL2/TaCBL3-TaCIPK29) were found to be responsive to distinct abiotic stresses. These results suggest that wheat CBL and CIPK genes may collectively mediate crosstalk of multiple stress signaling pathways through the formation of different CBL-CIPK complexes to decode stress-specific Ca2+signaling.
4) The roles of the TaCIPK14and TaCIPK29gene in response to various abiotic stresses were further investigated. TaCIPK14gene was upregulated under cold or when treated with salt, PEG or exogenous stresses related signaling molecules including ABA, ethylene and H2O2. Subcellular localization assay revealed the presence of TaCIPK14throughout the cell. Phenotype analysis showed that overexpression of TaCIPK14in tobacco enhanced cold and salt stress tolerance. Overexpression of TaCIPK14enhanced cold and salt stress tolerance by regulating the expression of antioxidant genes or stress related genes such as NtCAT, NtDREB3, NtLEA5, etc and enhancing the antioxidant system to reduce ROS accumulation and relieve membrane damage. Moreover, enhanced salt stress tolerance in TaCIPK14overexpressing plants was also attributed to decreased Na+content and elevated K+/Na+ratio.
5) TaCIPK29transcript was induced by NaCl, cold, methyl viologen (MV), ABA and ethylene treatments. Overexpression of TaCIPK29in tobacco resulted in increased salt tolerance. Further investigation showed that transgenic tobacco seedlings retained high K+/Na+ratio and Ca2+content by up-regulating some transporter genes such as NtSOSI, NtNHX2, NtNHX4and NtCAX3and reduced H2O2accumulation and membrane damage through enhancing the activities and expression of CAT and POD under salt stress. Finally, TaCIPK29was localized throughout cells and it interacted with TaCBL2, TaCBL3, NtCBL2, NtCBL3and NtCAT1. Taken together, our results showed that TaCIPK29functions as a positive factor under salt stress, involved in the regulation of ion and ROS homeostasis.
In conclusion, wheat CBL and CIPK genes were found to be abiotic stress responsive genes as those in rice and Arabidopsis. Different CBL or CIPK genes may differentially contribute to the plants' tolerance to distinct stresses. TaCIPK14and TaCIPK29gene may be useful candidate genes for developing stress-tolerant crops including wheat. Our work may facilitate further functional studies of these two important gene families and is beneficial for further molecular breeding in wheat.
1)小麦CBL和CIPK蛋白与水稻同源的CBL和CIPK蛋白的氨基酸序列和结构高度相似,这些CBL蛋白均含有四个能结合Ca2+的EF-hands结构域和一个FPSF结构域；CIPK蛋白都含有激酶结构域,FISL结构域和PPI结构域。进化分析显示,小麦CBL蛋白和CIPK蛋白和水稻同源CBL蛋白和CIPK蛋白都归类于相同的亚族。
2)基因表达分析结果表明,小麦特定的CBL或CIPK基因对冷、盐、渗透胁迫和ABA处理的响应模式不同,而且同一处理能够被不同的CBL或CIPK基因感应,这说明小麦CBL和CIPK基因可能介导对不同胁迫信号的交联过程。
3)利用酵母双杂交技术分析发现,小麦不同的CBL和CIPK蛋白之间的相互作用的特异性和强度不同。一些重要的特异性的CBL-CIPK互作子如：TaCBL1-TaCIPK9, TaCBL1-TaCIPK23, TaCBL2-TaCIPK2, TaCBL3-TaCIPK2, TaCBL2-TaCIPK29和TaCBL3-TaCIPK29可能参与小麦对不同非生物胁迫的响应。这些结果表明,小麦CBL和CIPK对不同非生物胁迫的信号转导是通过形成不同的CBL-CIPK复合物来解码特定胁迫下产生的特异Ca2+信号。
4)系统分析了TaCIPK14基因和TaCIPK29基因在非生物胁迫反应中的作用。TaCIPK14基因受冷、渗透胁迫和盐胁迫等非生物胁迫以及胁迫相关信号分子ABA、乙烯和H202诱导。证明了在烟草中过表达TaCIPK14基因可提高转基因烟草植株对冷和盐胁迫的抗性。TaCIPK14提高对冷和盐的抗性是通过调控一些抗氧化酶或胁迫相关基因如NtCAT, NtDREB3和NtLEA等的表达来实现的。增强的抗氧化酶的活性有助于清除逆境下产生的过量ROS,减轻ROS对细胞膜的损伤。同时,TaCIPK14能降低转基因烟草在盐胁迫下的Na+含量和提高细胞的K+/Na+比率,这也有助于植物对盐胁迫的抗性。
5) TaCIPK29基因能够被盐、冷、甲基紫晶(MV)、ABA和乙烯诱导。过表达TaCIPK29基因的烟草植株增强了对盐胁迫的抗性。TaCIPK29通过提高一些通道蛋白基因如NtSOS1, NtNHX2, NtNHX4和NtCAX3的表达来维持高的K+/Na+比和Ca2+含量以及通过增强CAT和POD的表达和活性来减少H2O2对细胞膜的损伤来抵抗盐胁迫的伤害。TaCIPK29没有特定的定位信号,在整个细胞中都有分布。。TaCIPK29不仅能够特异地与小麦的TaCBL2和TaCBL3互作,也能与烟草的NtCBL2和NtCBL3互作,说明TaCIPK29通过CBL2和CBL3传递Ca2+信号。同时TaCIPK29还能与烟草的NtCAT1相互作用,增强其活性,这有助于清除过多的H202。因此,TaCIPK29是一个盐胁迫中的正调控因子,参与了盐胁迫下对离子和ROS的调控。
综上所述,如同水稻和拟南芥中同源的CBL和CIPK基因一样,小麦CBL和CIPK基因也是非生物胁迫响应基因。不同的CBL和CIPK基因可能赋予植物对不同逆境的抗性。已鉴定的抗逆基因TaCIPK14和TaCIPK29可作为培育转基因抗逆小麦新品种的候选基因。本研究为深入研究小麦中这两个重要的基因家族的功能和分子机制奠定了基础,同时也有助于小麦抗逆分子育种研究与开发。
Wheat is a world staple crop, and its acreage, total production and trade ranks the first place among food crops. However, wheat production is often constrained by various abiotic stresses, such as drought, salinity, and extreme temperatures. Understanding the molecular mechanism of the abiotic stress responses can facilitate the genetic improvement of stress tolerance in wheat. Calcium, as a second messenger, is involved in the mediation of various responses to different environmental stresses and related to highly complex signal transduction pathways. Calcineurin B-like proteins (CBLs) and their target proteins, the CBL-interacting protein kinases (CIPKs), have emerged as key Ca2+-mediated signaling components in response to various abiotic stresses in many plants. However, the CBL and CIPK genes in wheat have not yet been comprehensively studied to date, except one CIPK gene (WPK4) which has been reported to mediate cytokine signaling transduction and the response to light and nutrient deprivation. In this study, seven CBL genes and eight CIPK genes were amplified from wheat genome using the RACE technique and their preferential interaction and differential responses to various abiotic stresses were investigated. The roles of two wheat CIPK genes TaCIPK14and TaCIPK29in response to abiotic stresses were further investigated. The main results are as follows:
1) Wheat CBLs and CIPKs were found to be similar to their counterparts in rice in motif structure and subgroup classification. These CBL proteins have four EF-hands and FPSF domains, while CIPK proteins contain kinase domain, FISL domain and PPI domain.
2) The isolated wheat CBL and CIPK genes were found to be expressed differentially in various tissues and in response to different abiotic stresses including cold, salt, and osmotic stresses and exposure to the phytohormone abscisic acid (ABA). Furthermore, we also found that one CBL or CIPK gene was able to respond to several treatments, and one treatment was sensed by multiple CBL or CIPK genes. Thus, TaCBLs and TaCIPKs were found to mediate crosstalk among different signaling pathways.
3) The preferential interactions of TaCBLs and TaCIPKs were identified using a yeast two-hybrid assay. Several important, specific CBL-CIPK interaction partners (TaCBL1-TaCIPK9, TaCBL1-TaCIPK23, TaCBL2-TaCIPK2, and TaCBL3-TaCIPK2, TaCBL2/TaCBL3-TaCIPK29) were found to be responsive to distinct abiotic stresses. These results suggest that wheat CBL and CIPK genes may collectively mediate crosstalk of multiple stress signaling pathways through the formation of different CBL-CIPK complexes to decode stress-specific Ca2+signaling.
4) The roles of the TaCIPK14and TaCIPK29gene in response to various abiotic stresses were further investigated. TaCIPK14gene was upregulated under cold or when treated with salt, PEG or exogenous stresses related signaling molecules including ABA, ethylene and H2O2. Subcellular localization assay revealed the presence of TaCIPK14throughout the cell. Phenotype analysis showed that overexpression of TaCIPK14in tobacco enhanced cold and salt stress tolerance. Overexpression of TaCIPK14enhanced cold and salt stress tolerance by regulating the expression of antioxidant genes or stress related genes such as NtCAT, NtDREB3, NtLEA5, etc and enhancing the antioxidant system to reduce ROS accumulation and relieve membrane damage. Moreover, enhanced salt stress tolerance in TaCIPK14overexpressing plants was also attributed to decreased Na+content and elevated K+/Na+ratio.
5) TaCIPK29transcript was induced by NaCl, cold, methyl viologen (MV), ABA and ethylene treatments. Overexpression of TaCIPK29in tobacco resulted in increased salt tolerance. Further investigation showed that transgenic tobacco seedlings retained high K+/Na+ratio and Ca2+content by up-regulating some transporter genes such as NtSOSI, NtNHX2, NtNHX4and NtCAX3and reduced H2O2accumulation and membrane damage through enhancing the activities and expression of CAT and POD under salt stress. Finally, TaCIPK29was localized throughout cells and it interacted with TaCBL2, TaCBL3, NtCBL2, NtCBL3and NtCAT1. Taken together, our results showed that TaCIPK29functions as a positive factor under salt stress, involved in the regulation of ion and ROS homeostasis.
In conclusion, wheat CBL and CIPK genes were found to be abiotic stress responsive genes as those in rice and Arabidopsis. Different CBL or CIPK genes may differentially contribute to the plants' tolerance to distinct stresses. TaCIPK14and TaCIPK29gene may be useful candidate genes for developing stress-tolerant crops including wheat. Our work may facilitate further functional studies of these two important gene families and is beneficial for further molecular breeding in wheat.
引文
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