Na_2CO_3胁迫下甜高粱CBL基因家族的表达模式分析及功能初探
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摘要
土壤盐碱化给作物的生长带来了严重的危害,并对农业的发展产生了不利的影响。由于当今世界能源安全和环境问题的凸显,生物质能源的开发已迫在眉睫。甜高粱具有很多优良的生物学特性,使之成为最具潜力的生物质能源作物之一。为了了解甜高粱耐盐碱的生理特性以及发掘CBL基因家族中响应盐碱胁迫的功能基因以用于甜高粱耐盐碱新品种的遗传改良,本研究选取甜高粱“能饲Ⅰ号”品种为实验材料,分析了在Na2CO3胁迫条件下的生理响应,确定了一个适宜的盐碱胁迫处理浓度;通过实时荧光定量技术分析了盐碱胁迫条件下基因的表达模式及在正常生长条件下不同组织不同发育时期的表达模式;通过Gateway技术克隆了盐碱胁迫响应基因并转化拟南芥;通过酵母双杂交技术分析蛋白间的互作;同时,通过生物信息学的方法分析了该基因家族的生物信息学特性,取得的结果如下:
(1)通过生理实验确定了活力指数,胚根和胚芽长度可以作为甜高粱耐盐碱品种的筛选的指标。同时,确定了甜高粱幼苗期Na2CO3胁迫处理的适宜浓度:100mM。
(2)通过实时荧光定量的方法分析了甜高粱CBL基因家族在不同发育时期不同组织中的表达特性及幼苗期SbCBL基因家族对Na2CO3胁迫的响应模式,结果发现:
该家族基因的表达具有组织特异性,SbCBL06、SbCBL08 mRNA相对表达水平在幼叶中最高,而其他6个SbCBL基因mRNA相对表达水平均在成熟叶片中最高;
甜高粱CBL基因家族对Na2CO3胁迫响应呈现出不同的表达模式,SbCBL01、SbCBL07主要在幼苗根部呈现持续的上调表达,SbCBL06主要在幼苗叶片中响应。
(3)应用生物信息学的方法对SbCBL基因家族进行生物信息学分析。
首先通过数据库EST比对发现大部分SbCBL均有与之对应的受逆境胁迫诱导的EST序列,因此初步推测,该家族基因在抵抗逆境方面可能具有重要功能。
SbCBL家族氨基酸序列多重比对及EF基序在线查找发现,除了SbCBL08含有2个EF-hand,其它SbCBL中均含有3个典型的EF-hand保守结构域,并且在SbCBL01、SbCBL04、SbCBL05和SbCBL08中具有N-豆蔻酰化结构域(MGXXXS/T)。
对SbCBL家族基因上游5′-UTR 2.5kb序列进行预测分析,结果表明该家族基因启动子区均含有多种与逆境胁迫及植物激素相关的顺式作用元件。
通过分析SbCBL基因家族外显子—内含子结构,发现CBL基因家族内含子数目在物种间及物种内均含有一定的保守性。
通过进化树分析了SbCBL基因家族分别与拟南芥、水稻、玉米、葡萄、杨树和绿藻等物种中的CBL基因家族在进化上的亲缘关系。结果发现,在这几个物种中,SbCBL基因家族与水稻中的亲缘关系最近。
(4)通过Gateway技术克隆了SbCBL01基因,构建到植物表达载体pLeela,利用农杆菌介导的沾花侵染的方法转化拟南芥,得到6个T1代的转基因株系;
(5)将SbCBL01及多个能够响应Na2CO3胁迫的SbCIPK基因构建到pDEST22及pDEST32载体共转化酵母细胞,结果发现, SbCBL01蛋白可能与SbCIPK24及SbCIPK30蛋白互作;
Saline-alkali soil contains excessive sodium carbonate (Na2CO3) and sodium bicarbonate (NaHCO3), which inhibits most plant growth and development. The tradition fossil fuel is getting shortage and leads to a lot of environment problems, The development of renewable energy sources is urgently needed. Sweet sorghum (Sorghum bicolor L.) is characterized by drought and flood resistance, salt tolerance and high efficiency of biomass accumulation. It is considered to be one of the most promising bio-energy crops. In order to understand the function and the physiological expression pattern of SbCBL gene family in sweet sorghum under sodium carbonate stress and utilize this experimental results to obtain new sorghum cultivar which can tolerate sodium carbonate stress, we use sweet sorghum (Sorghum bicolor L.) cv. NengSiⅠto be our materials. In our research, we analyzed the physiological response of sweet sorghum, and identified a eligible sodium carbonate concentration. We used real-time quantitative polymerase chain reaction (RTqPCR) to analyze the expression pattern of SbCBL gene family under sodium carbonate stress and the expression pattern of SbCBL gene family in different tissues under regular condition. We cloned special gene by Gateway technology and introduced it into Arabidopsis. We analyzed the protein-protein interaction by yeast two hybrid. Meanwhile, through the bioinformatics analysis, we identified the character of SbCBL gene family. The results are as follows:
(1) The results of physiological experiment showed that vitality index, radicle and the length of plumule can be the index of sorghum cultivar which can tolerate sodium carbonate stress, and we identified a eligible sodium carbonate treatment concentration: 100mM Na_2CO_3.
(2) We used real-time quantitative polymerase chain reaction to analyze the expression pattern of SbCBL gene family under sodium carbonate stress and the expression pattern of SbCBL gene family in different tissues under normal condition. The results showed that SbCBL genes in sweet sorghum have different tissue-specific expression patterns under normal growth condition, the transcription of SbCBL06 and SbCBL08 were most abundant in leaves at seedling stage and others were higher expression in matured leaves. In addition, SbCBL genes showed various sodium carbonate stress responsive patterns in sweet sorghum seedlings treated with sodium carbonate, the transcriptional level of SbCBL01 and SbCBL07 showed sustained increase in roots and a similar pattern appeared in the transcription of SbCBL06 in shoots.
(3) Bioinformatic analysis of SbCBLs:
Firstly, Each SbCBL had its homologous ESTs in the database. Some of these ESTs were related to abiotic stresses. The results showed that SbCBL gene family might play an important role in abiotic stress.
Secondly, Multiple alignments revealed that sequences of SbCBLs are highly conserved and most SbCBLs have three typical EF-hands structure. Four SbCBL proteins (SbCBL01, SbCBL04, SbCBL05, SbCBL08) have a conserved N-myristoylation domain (MGXXXS/T).
Thirdly, Many stress responsive and phytohormones responsive cis-elements are found in the promoter region of SbCBLs. The conservation in intron number of the same or different species infers that the introns may have some effect on the function of CBL gene family.
Phylogenetic analysis of SbCBL gene family between sweet sorghum and other plants (Arabidopsis, rice, maize, Vitis vinifera, poplar and chlorella ) suggested that sweet sorghum CBL gene family had the most closely genetic relationship with rice.
(4) We cloned SbCBL01 by Gateway technology and introduced it into the expression vector pLeela. We used the floral dip method of Agrobacterium-mediated transformation to infect Arabidopsis. SbCBL01 has been inserted into the genomic DNA of Arabidopsis. We obtained six T1 transgenic lines .
(5) We constructed SbCBL01 and potential target SbCIPK gene into pDEST22 and pDEST32, respectively and analyzed SbCBL and SbCIPK interaction. The result showed that SbCBL01 might interact with SbCIPK24 and SbCIPK30.
(1)通过生理实验确定了活力指数,胚根和胚芽长度可以作为甜高粱耐盐碱品种的筛选的指标。同时,确定了甜高粱幼苗期Na2CO3胁迫处理的适宜浓度:100mM。
(2)通过实时荧光定量的方法分析了甜高粱CBL基因家族在不同发育时期不同组织中的表达特性及幼苗期SbCBL基因家族对Na2CO3胁迫的响应模式,结果发现:
该家族基因的表达具有组织特异性,SbCBL06、SbCBL08 mRNA相对表达水平在幼叶中最高,而其他6个SbCBL基因mRNA相对表达水平均在成熟叶片中最高;
甜高粱CBL基因家族对Na2CO3胁迫响应呈现出不同的表达模式,SbCBL01、SbCBL07主要在幼苗根部呈现持续的上调表达,SbCBL06主要在幼苗叶片中响应。
(3)应用生物信息学的方法对SbCBL基因家族进行生物信息学分析。
首先通过数据库EST比对发现大部分SbCBL均有与之对应的受逆境胁迫诱导的EST序列,因此初步推测,该家族基因在抵抗逆境方面可能具有重要功能。
SbCBL家族氨基酸序列多重比对及EF基序在线查找发现,除了SbCBL08含有2个EF-hand,其它SbCBL中均含有3个典型的EF-hand保守结构域,并且在SbCBL01、SbCBL04、SbCBL05和SbCBL08中具有N-豆蔻酰化结构域(MGXXXS/T)。
对SbCBL家族基因上游5′-UTR 2.5kb序列进行预测分析,结果表明该家族基因启动子区均含有多种与逆境胁迫及植物激素相关的顺式作用元件。
通过分析SbCBL基因家族外显子—内含子结构,发现CBL基因家族内含子数目在物种间及物种内均含有一定的保守性。
通过进化树分析了SbCBL基因家族分别与拟南芥、水稻、玉米、葡萄、杨树和绿藻等物种中的CBL基因家族在进化上的亲缘关系。结果发现,在这几个物种中,SbCBL基因家族与水稻中的亲缘关系最近。
(4)通过Gateway技术克隆了SbCBL01基因,构建到植物表达载体pLeela,利用农杆菌介导的沾花侵染的方法转化拟南芥,得到6个T1代的转基因株系;
(5)将SbCBL01及多个能够响应Na2CO3胁迫的SbCIPK基因构建到pDEST22及pDEST32载体共转化酵母细胞,结果发现, SbCBL01蛋白可能与SbCIPK24及SbCIPK30蛋白互作;
Saline-alkali soil contains excessive sodium carbonate (Na2CO3) and sodium bicarbonate (NaHCO3), which inhibits most plant growth and development. The tradition fossil fuel is getting shortage and leads to a lot of environment problems, The development of renewable energy sources is urgently needed. Sweet sorghum (Sorghum bicolor L.) is characterized by drought and flood resistance, salt tolerance and high efficiency of biomass accumulation. It is considered to be one of the most promising bio-energy crops. In order to understand the function and the physiological expression pattern of SbCBL gene family in sweet sorghum under sodium carbonate stress and utilize this experimental results to obtain new sorghum cultivar which can tolerate sodium carbonate stress, we use sweet sorghum (Sorghum bicolor L.) cv. NengSiⅠto be our materials. In our research, we analyzed the physiological response of sweet sorghum, and identified a eligible sodium carbonate concentration. We used real-time quantitative polymerase chain reaction (RTqPCR) to analyze the expression pattern of SbCBL gene family under sodium carbonate stress and the expression pattern of SbCBL gene family in different tissues under regular condition. We cloned special gene by Gateway technology and introduced it into Arabidopsis. We analyzed the protein-protein interaction by yeast two hybrid. Meanwhile, through the bioinformatics analysis, we identified the character of SbCBL gene family. The results are as follows:
(1) The results of physiological experiment showed that vitality index, radicle and the length of plumule can be the index of sorghum cultivar which can tolerate sodium carbonate stress, and we identified a eligible sodium carbonate treatment concentration: 100mM Na_2CO_3.
(2) We used real-time quantitative polymerase chain reaction to analyze the expression pattern of SbCBL gene family under sodium carbonate stress and the expression pattern of SbCBL gene family in different tissues under normal condition. The results showed that SbCBL genes in sweet sorghum have different tissue-specific expression patterns under normal growth condition, the transcription of SbCBL06 and SbCBL08 were most abundant in leaves at seedling stage and others were higher expression in matured leaves. In addition, SbCBL genes showed various sodium carbonate stress responsive patterns in sweet sorghum seedlings treated with sodium carbonate, the transcriptional level of SbCBL01 and SbCBL07 showed sustained increase in roots and a similar pattern appeared in the transcription of SbCBL06 in shoots.
(3) Bioinformatic analysis of SbCBLs:
Firstly, Each SbCBL had its homologous ESTs in the database. Some of these ESTs were related to abiotic stresses. The results showed that SbCBL gene family might play an important role in abiotic stress.
Secondly, Multiple alignments revealed that sequences of SbCBLs are highly conserved and most SbCBLs have three typical EF-hands structure. Four SbCBL proteins (SbCBL01, SbCBL04, SbCBL05, SbCBL08) have a conserved N-myristoylation domain (MGXXXS/T).
Thirdly, Many stress responsive and phytohormones responsive cis-elements are found in the promoter region of SbCBLs. The conservation in intron number of the same or different species infers that the introns may have some effect on the function of CBL gene family.
Phylogenetic analysis of SbCBL gene family between sweet sorghum and other plants (Arabidopsis, rice, maize, Vitis vinifera, poplar and chlorella ) suggested that sweet sorghum CBL gene family had the most closely genetic relationship with rice.
(4) We cloned SbCBL01 by Gateway technology and introduced it into the expression vector pLeela. We used the floral dip method of Agrobacterium-mediated transformation to infect Arabidopsis. SbCBL01 has been inserted into the genomic DNA of Arabidopsis. We obtained six T1 transgenic lines .
(5) We constructed SbCBL01 and potential target SbCIPK gene into pDEST22 and pDEST32, respectively and analyzed SbCBL and SbCIPK interaction. The result showed that SbCBL01 might interact with SbCIPK24 and SbCIPK30.
引文
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