玉米幼苗四个水份胁迫响应基因在不同耐旱自交系中的表达分析
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摘要
为了深入研究玉米耐旱的分子机理,本研究用一耐旱玉米自交系“81565”,构建了水份胁迫下玉米幼苗差异表达mRNA的cDNA文库,并筛选鉴定了其中5个EST序列。其中两个是玉米的类胰蛋白丝氨酸蛋白酶基因(tryspin-like serineproteases,Dege)和不依赖辅助因子的磷酸甘油酸变位酶基因(cofactor-independent phosphoglycerate mutase,PGAM-i),一个基因编码具有NADH脱氢酶活性的还原酶,两个功能未知,这里命名为E4和F4。类胰蛋白丝氨酸蛋白酶(DegP)广泛参与生物耐热激反应和高等植物中的光抑制修复反应。拟南芥物种的AtDegP1,AtDegP5,AtDegP8和AtDegP2蛋白通过降解氧化变性的D1蛋白,而恢复PSⅡ的光合作用。耐盐植物冰草(Mesembryanthemu crystallinum)中,磷酸甘油酸变位酶基因(PGAM-i)的表达响应多种非生物逆境胁迫。基因E4的编码蛋白是含有DUF6和TPT结构域的膜整合蛋白。为了研究DegP,PGAM-i,E4和F4这4个基因如何响应水份胁迫,本实验用实时定量PCR技术对4个基因在水份胁迫下(20%PEG处理)的6个玉米自交系中的表达进行了分析。6个玉米自交系“81565”,“87-1”,“丹340”,“R09”,“200B”,“ES40”具有不同的田间耐旱性。DegP基因在植物的抗热激和PSⅡ光合抑制修复中起到重要作用,为近一步研究此基因在玉米耐旱中的作用和对产量的影响,克隆了玉米的DegP基因,并作了初步的生物信息学分析。主要的研究结果如下:
1.用耐旱自交系“81565”和抑制差减杂交,反式Northern技术分离鉴定了玉米幼苗水份胁迫条件下上调表达的5个EST序列,其中三个分别是编码DegP,PGAM-i,NADH还原酶的基因,两个功能未知。
2.自交系“81565”中Dege,PGAM-i,E4和F4基因的表达响应水份胁迫。正常生长条件下,DegP,PGAM-i,E4和F4基因都有一定量的表达。在PEG处理的6h时,水份胁迫抑制了4个基因的表达,它们的表达量都低于胁迫处理前的表达量。随着胁迫处理时间的延长,4基因的表达量开始上升。在胁迫12 h时,PGAM-i基因的相对表达量达到了处理前的1.4倍。在胁迫24 h时,DegP基因的相对表达量是处理前的0.6倍;E4基因是处理前的1.2倍,F4基因是处理前的1.6倍。水份胁迫处理下,PGAM-i,E4和F4基因的表达都上调了。
3.水份胁迫下,6玉米自交系中4基因的表达模式有相似和不同。耐旱自交系“87-1”和“R09”4基因的表达模式和“81565”自交系的相似。耐旱自交系“200B”中DegP基因的表达受到抑制,水份胁迫24 h时,PGAM-i基因的表达量被诱导到处理前的6倍,E4被诱导到2.6倍,F4基因被诱导到2倍。不耐旱自交系“丹340”中DegP和E4基因被下调表达,PGAM-i基因的表达在胁迫24 h时得到恢复,F4基因受胁迫诱导上调表达。不耐旱自交系“ES40”4基因的表达都受水份胁迫抑制,表达量一直降低。这从分子水平也反映出“ES40”自交系的弱耐旱性。
4.克隆了玉米自交系“87-1”中的DegP基因,发现其存在缺失突变,并分析了DegP基因此位点的序列多样性。生物信息学分析发现玉米基因DegP和拟南芥的AtDegP8基因同源性非常高,二者间氨基酸序列的相似性是66%。分析同时发现玉米基因组中存在一个和AtDegP1基因同源性较高的DegP家族的另一成员,二者N端氨基酸序列比较,其一致性是53%。
本文的研究结果表明,玉米中DegP,PGAM-i,E4,F4基因的表达响应水份胁迫,它们的表达对增加玉米的耐旱性有着重要作用。6个自交系间的比较分析显示了它们在干旱胁迫下对4个基因的不同调控模式。其中三个耐旱自交系“81565”,“R09”,“87-1”对4个基因的上调表达显示了其在水分胁迫时,它们更迅速的调控能力。自交系“ES40”在水份胁迫下4个基因的表达一直受到抑制,这和它的弱耐旱性一致。
高等植物的DegP基因家族广泛参与了植物的耐热激反应和光抑制修复反应。由于玉米DegP基因和AtDegP8基因的蛋白序列相似性很高,推测玉米的此基因为AtDegP8的同源基因。拟南芥的AtDegP1,AtDegP5,AtDegP8蛋白位于叶绿体内囊体内腔中,它们和位于内囊体基质的AtDegP2蛋白,及FtsH蛋白一起修复光抑制反应,恢复光合反应正常进行。这些结果显示DegP基因家族对提高玉米物种的耐旱性有重要作用,它们对逆境胁迫下的产量也有重要影响。
玉米DegP基因存在不同基因型,如自交系“87-1”中的缺失突变型,推测其不能合成有功能的完整DegP蛋白。这种基因型和自交系耐旱能力的关系,有待近一步研究。
To further understand how maize responds to drought stress,we constructed the cDNA library that maize seedling specially responds to water stress utilizing one drought-tolerant inbred "81565",and have identified 5 EST sequence.After the blast analysis,we identified the characters of three of them.One is the tryspin-like serine proteases gene(DegP),others are the cofactor-independent phosphoglycerate mutase (PGAM-i) and the gene encoding the G subunit of NADH dehydrogenase complex. Other two genes were designated as E4 and F4,without clear function.Tryspin-like serine proteases(DegP) widely involved in heat stress response of eukaryote and prokaryote,also take part in repair action against photoinhibition of higher plants.The AtDegP1,AtDegP5,AtDegP8 and AtDegP2 enzymes in Arabidopsis degrade the photodamaged D1 protein to rescue the PSⅡphotosynthesis.One facultative halophyte,its name is ice plant(Mesembryanthemu crystallinum).The PGAM-i gene of this plant,which transcriptional level is induced by several abiotic stress.The encoding protein of E4 gene is a membrane integration protein with DUF6 and TPT domain.To disclose how the 4 maize genes,DegP,PGAM-i,E4 and F4 response water stress,we employed real time quantitative PCR technology and analyzed 4 genes expression profile of 6 maize inbred lines under water stress(20%PEG treatment).The 6 maize inbred lines "81565","87-1","R09","200B","Dan340" and "ES40" have different drought-tolerant ability in field surrounding.DegP gene has been identified that takes part in heat shock response and repair action against PSⅡphotoinhibition of higher plants.For finding out the function of DegP gene in maize drought-tolerance,we cloned the maize DegP gene.Based on the DegP gene sequence,we analyzed the primary character of DegP gene using biological information method.Our results are mainly as follows:
1.Utilizing the drought-tolerant maize inbreed "81565",the techniques of Suppression subtractive hybridization(SSH) and reverse Northern hybridization,we isolated and identified 5 up-regulated EST sequence from the water stressed maize seedlings.Three of them are the genes encoding DegP,PGAM-i,and the G subunit protein of NADH dehydrogenase complex,respectively.The functions of others are not identified.
2.The expression of DegP,PGAM-i,E4 and F4 genes respond to water deficit in drought-tolerant inbred "81565".DegP,PGAM-i,E4 and F4 genes express certain level under normal condition.After 6 h of PEG treatment,water stress repressed the expression of 4 genes.Their expressional level all decreased compared with the level of control treatment.But the expressional level up-regulated with the treatment time elongation.PGAM-i gene expressional level was 1.4 times compared with control level after stress treatment 12 h.DegP gene expression increased to 0.6 times after stress treatment 24 h.The expressional level of E4 and F4 genes were up-regulated and were 1.2 times and 1.6 times compared with control level after 24 hours, respectively.
3.The 4 genes expression profiles of 6 maize inbred lines under water stress have similarity and difference.The 4 genes expression profiles of drought-tolerant lines "87-1" and "R09" were similar with "81565" line.The expression of DegP gene of drought-tolerant line "200B" was repressed by water stress,but after stress 24 h, the expressional level of PGAM-i was induced and was 6 times compared with the control level,the expression level of E4 and F4 genes were induced to 2.6 and 2 times compared with the control level,respectively.The DegP and E4 genes expression of Drought-sensitive "Dan340" were down-regulated.After 24 h treatment,the PGAM-i gene expression level recovered,F4 gene expression was induced to high level.The 4 genes expression in another drought-sensitive inbred line "ES40" were all repressed, the expression level decreased along the treatment time.The sign also indicated the drought-sensitive character of"ES40" line in molecular level.
4.This research based on the EST sequence of DegP,cloned the DegP gene from the inbred 87-1.We have identified the deletion mutation occurred in the 87-1 DegP gene,and analyzed the sequence diversity at the mutation site.Alignment analysis identified the DegP gene is the homolog of Arabidopsis AtDegP8.the identity of amino acid sequence between them is 66%.Other member of DegP gene family was also found in maize genome,which amino acid sequence was more similar with Arabidopsis AtDegP1 gene,the N terminal amino acid sequences of them were aligned,the identity is 53%.
The research demonstrated that DegP,PGAM-i,E4 and F4 gene play a role in maize drought response,and their expression level contribute to the plant tolerance to water stress.The comparative analysis between 6 inbred lines revealed the different regulation model for the 4 genes.The 4 genes up-regulation in 3 drought-tolerant lines "81565","R09","87-1" indicates the 3 inbred lines have rapid regulation ability.The 4 genes up-regulation in line "ES40" were repressed along the treatment time,this is accordant with the drought-sensitive trait of"ES40".
The DegP family genes in higher plants contribute to the acquirement to adapting abiotic stress.The high identity of amino acid sequence between maize DegP and AtDegP8 suggest that the maize DegP is the ortholog of AtDegP8.The AtDegP1, AtDegP5,AtDegP8 proteins locate in the lumen of thylakoid,the AtDegP2 protein locates in stroma.The four proteins and the FtsH protein together eliminate the damage by photoinhibition,and recover to the normal photosynthesis activity.These experiments showed that DegP gene family play important role in maize drought tolerance,and in maize production under water stress.
The maize DegP gene have different genotype,one is the deletion mutation allele in inbred line "87-1",which presumed that it could not synthesis the intact structural and functional DegP protein.The relationship between this genotype and the ability of drought tolerance in different lines is still deciphered.
1.用耐旱自交系“81565”和抑制差减杂交,反式Northern技术分离鉴定了玉米幼苗水份胁迫条件下上调表达的5个EST序列,其中三个分别是编码DegP,PGAM-i,NADH还原酶的基因,两个功能未知。
2.自交系“81565”中Dege,PGAM-i,E4和F4基因的表达响应水份胁迫。正常生长条件下,DegP,PGAM-i,E4和F4基因都有一定量的表达。在PEG处理的6h时,水份胁迫抑制了4个基因的表达,它们的表达量都低于胁迫处理前的表达量。随着胁迫处理时间的延长,4基因的表达量开始上升。在胁迫12 h时,PGAM-i基因的相对表达量达到了处理前的1.4倍。在胁迫24 h时,DegP基因的相对表达量是处理前的0.6倍;E4基因是处理前的1.2倍,F4基因是处理前的1.6倍。水份胁迫处理下,PGAM-i,E4和F4基因的表达都上调了。
3.水份胁迫下,6玉米自交系中4基因的表达模式有相似和不同。耐旱自交系“87-1”和“R09”4基因的表达模式和“81565”自交系的相似。耐旱自交系“200B”中DegP基因的表达受到抑制,水份胁迫24 h时,PGAM-i基因的表达量被诱导到处理前的6倍,E4被诱导到2.6倍,F4基因被诱导到2倍。不耐旱自交系“丹340”中DegP和E4基因被下调表达,PGAM-i基因的表达在胁迫24 h时得到恢复,F4基因受胁迫诱导上调表达。不耐旱自交系“ES40”4基因的表达都受水份胁迫抑制,表达量一直降低。这从分子水平也反映出“ES40”自交系的弱耐旱性。
4.克隆了玉米自交系“87-1”中的DegP基因,发现其存在缺失突变,并分析了DegP基因此位点的序列多样性。生物信息学分析发现玉米基因DegP和拟南芥的AtDegP8基因同源性非常高,二者间氨基酸序列的相似性是66%。分析同时发现玉米基因组中存在一个和AtDegP1基因同源性较高的DegP家族的另一成员,二者N端氨基酸序列比较,其一致性是53%。
本文的研究结果表明,玉米中DegP,PGAM-i,E4,F4基因的表达响应水份胁迫,它们的表达对增加玉米的耐旱性有着重要作用。6个自交系间的比较分析显示了它们在干旱胁迫下对4个基因的不同调控模式。其中三个耐旱自交系“81565”,“R09”,“87-1”对4个基因的上调表达显示了其在水分胁迫时,它们更迅速的调控能力。自交系“ES40”在水份胁迫下4个基因的表达一直受到抑制,这和它的弱耐旱性一致。
高等植物的DegP基因家族广泛参与了植物的耐热激反应和光抑制修复反应。由于玉米DegP基因和AtDegP8基因的蛋白序列相似性很高,推测玉米的此基因为AtDegP8的同源基因。拟南芥的AtDegP1,AtDegP5,AtDegP8蛋白位于叶绿体内囊体内腔中,它们和位于内囊体基质的AtDegP2蛋白,及FtsH蛋白一起修复光抑制反应,恢复光合反应正常进行。这些结果显示DegP基因家族对提高玉米物种的耐旱性有重要作用,它们对逆境胁迫下的产量也有重要影响。
玉米DegP基因存在不同基因型,如自交系“87-1”中的缺失突变型,推测其不能合成有功能的完整DegP蛋白。这种基因型和自交系耐旱能力的关系,有待近一步研究。
To further understand how maize responds to drought stress,we constructed the cDNA library that maize seedling specially responds to water stress utilizing one drought-tolerant inbred "81565",and have identified 5 EST sequence.After the blast analysis,we identified the characters of three of them.One is the tryspin-like serine proteases gene(DegP),others are the cofactor-independent phosphoglycerate mutase (PGAM-i) and the gene encoding the G subunit of NADH dehydrogenase complex. Other two genes were designated as E4 and F4,without clear function.Tryspin-like serine proteases(DegP) widely involved in heat stress response of eukaryote and prokaryote,also take part in repair action against photoinhibition of higher plants.The AtDegP1,AtDegP5,AtDegP8 and AtDegP2 enzymes in Arabidopsis degrade the photodamaged D1 protein to rescue the PSⅡphotosynthesis.One facultative halophyte,its name is ice plant(Mesembryanthemu crystallinum).The PGAM-i gene of this plant,which transcriptional level is induced by several abiotic stress.The encoding protein of E4 gene is a membrane integration protein with DUF6 and TPT domain.To disclose how the 4 maize genes,DegP,PGAM-i,E4 and F4 response water stress,we employed real time quantitative PCR technology and analyzed 4 genes expression profile of 6 maize inbred lines under water stress(20%PEG treatment).The 6 maize inbred lines "81565","87-1","R09","200B","Dan340" and "ES40" have different drought-tolerant ability in field surrounding.DegP gene has been identified that takes part in heat shock response and repair action against PSⅡphotoinhibition of higher plants.For finding out the function of DegP gene in maize drought-tolerance,we cloned the maize DegP gene.Based on the DegP gene sequence,we analyzed the primary character of DegP gene using biological information method.Our results are mainly as follows:
1.Utilizing the drought-tolerant maize inbreed "81565",the techniques of Suppression subtractive hybridization(SSH) and reverse Northern hybridization,we isolated and identified 5 up-regulated EST sequence from the water stressed maize seedlings.Three of them are the genes encoding DegP,PGAM-i,and the G subunit protein of NADH dehydrogenase complex,respectively.The functions of others are not identified.
2.The expression of DegP,PGAM-i,E4 and F4 genes respond to water deficit in drought-tolerant inbred "81565".DegP,PGAM-i,E4 and F4 genes express certain level under normal condition.After 6 h of PEG treatment,water stress repressed the expression of 4 genes.Their expressional level all decreased compared with the level of control treatment.But the expressional level up-regulated with the treatment time elongation.PGAM-i gene expressional level was 1.4 times compared with control level after stress treatment 12 h.DegP gene expression increased to 0.6 times after stress treatment 24 h.The expressional level of E4 and F4 genes were up-regulated and were 1.2 times and 1.6 times compared with control level after 24 hours, respectively.
3.The 4 genes expression profiles of 6 maize inbred lines under water stress have similarity and difference.The 4 genes expression profiles of drought-tolerant lines "87-1" and "R09" were similar with "81565" line.The expression of DegP gene of drought-tolerant line "200B" was repressed by water stress,but after stress 24 h, the expressional level of PGAM-i was induced and was 6 times compared with the control level,the expression level of E4 and F4 genes were induced to 2.6 and 2 times compared with the control level,respectively.The DegP and E4 genes expression of Drought-sensitive "Dan340" were down-regulated.After 24 h treatment,the PGAM-i gene expression level recovered,F4 gene expression was induced to high level.The 4 genes expression in another drought-sensitive inbred line "ES40" were all repressed, the expression level decreased along the treatment time.The sign also indicated the drought-sensitive character of"ES40" line in molecular level.
4.This research based on the EST sequence of DegP,cloned the DegP gene from the inbred 87-1.We have identified the deletion mutation occurred in the 87-1 DegP gene,and analyzed the sequence diversity at the mutation site.Alignment analysis identified the DegP gene is the homolog of Arabidopsis AtDegP8.the identity of amino acid sequence between them is 66%.Other member of DegP gene family was also found in maize genome,which amino acid sequence was more similar with Arabidopsis AtDegP1 gene,the N terminal amino acid sequences of them were aligned,the identity is 53%.
The research demonstrated that DegP,PGAM-i,E4 and F4 gene play a role in maize drought response,and their expression level contribute to the plant tolerance to water stress.The comparative analysis between 6 inbred lines revealed the different regulation model for the 4 genes.The 4 genes up-regulation in 3 drought-tolerant lines "81565","R09","87-1" indicates the 3 inbred lines have rapid regulation ability.The 4 genes up-regulation in line "ES40" were repressed along the treatment time,this is accordant with the drought-sensitive trait of"ES40".
The DegP family genes in higher plants contribute to the acquirement to adapting abiotic stress.The high identity of amino acid sequence between maize DegP and AtDegP8 suggest that the maize DegP is the ortholog of AtDegP8.The AtDegP1, AtDegP5,AtDegP8 proteins locate in the lumen of thylakoid,the AtDegP2 protein locates in stroma.The four proteins and the FtsH protein together eliminate the damage by photoinhibition,and recover to the normal photosynthesis activity.These experiments showed that DegP gene family play important role in maize drought tolerance,and in maize production under water stress.
The maize DegP gene have different genotype,one is the deletion mutation allele in inbred line "87-1",which presumed that it could not synthesis the intact structural and functional DegP protein.The relationship between this genotype and the ability of drought tolerance in different lines is still deciphered.
引文
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