东南景天锌超积累过程中基因表达差异的cDNA-AFLP分析
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摘要
由于重金属超积累植物在治理重金属污染土壤方面,有广阔的应用前景,已引起国际土壤——植物营养学界和环境科学界的浓厚情趣和政府部门的高度重视。但目前对植物超积累重金属机制认识的不足限制了该技术的应用和改良。东南景天(Sedum alfredii Hance)是在我国东南部地区一些古老的铅锌矿区发现的一种新的锌/镉超积累植物,同时对铅具有富集作用。本研究以超积累东南景天为材料,以其他近缘种为对照,分析了铅锌矿区东南景天在长期的重金属胁迫条件下,进化成超积累植物后的基因组变异情况;以及利用cDNA-AFLP技术对超积累生态型东南景天在锌超积累过程中的基因表达差异进行研究;另外还采用水培方法,观察超积景天在锌镉积累过程中的GSH、H_2O_2、总硫含量随锌镉积累量的变化,探讨了GSH和H_2O_2在超积累过程中的作用,取得的主要研究结果如下:
1.植物长期生长在重金属污染的生境中,逐渐进化成不同的生态型,利用我国原生的Zn/Cd超积累生态型东南景天(Hyperaccumulated ecotype:HE)为材料,以与其亲缘关系较近的非超积累生态型东南景天(NHE)、圆叶景天(Sedum MakinoiMaxim:SMM)、凹叶景天(Sedum emarginatum Migo:SEM)、垂盆草(SedumSarmentoum Bunge:SSB)以及株芽景天(Sedum Bulbiferum Makino:SBM)为对照,利用RAPD方法,研究超积累生态型东南景天基因组变异的程度,并对部分超积累景天特有的PCR条带进行测序,分析特异基因片段与超积累的关系。遗传进化树分析表明:六种不同生态型的景天可以分成两组,其中两种东南景天为一组,另外四种为一组;超积累生态型和野生型东南景天有着相似的遗传背景,相似系数显示变异程度已超过种的界限。说明污染土壤对植物的进化有着非常大的影响;部分超积累景天特异PCR片段测序表明,ATP-硫酰化酶(ATP sulfuryase)、脱水调节相关蛋白及一些未知基因片段可能与超积累性状有一定的关系。
2.植物对重金属锌/镉超积累与过氧化氢产生关系的研究表明,在1000uM Zn~(2+)或200uM Cd~(2+)处理条件下,超积累生态型东南景天地上部Zn~(2+)的积累量高于Cd~(2+)的积累量。地上部Zn~(2+)/Cd~(2+)积累时间动力学显示在前15天,呈线形增加,随后积累速度下降。同样,地上部总硫、GSH以及H_2O_2的含量也是在前15天迅速增加,后10天逐渐下降。总硫和GSH的含量都是Cd~(2+)处理下较Zn~(2+)处理的高,而H_2O_2的含量则是Cd~(2+)处理下较Zn~(2+)处理的低;地上部Zn~(2+)含量变化与地上部GSH、总硫及H_2O_2变化趋势相似,而Cd~(2+)则是在第5天时含量就达到最大,以后开始逐渐下降。以上结果推测:含硫有机物可能不是超积累生态型东南景天Zn~(2+)/Cd~(2+)积累的主要配体,GSH的功能主要是解除体内游离态Zn~(2+)/Cd~(2+)的毒性,H_2O_2很可能是与超积累相关的一种信号分子。增加硫元素的供应对提高zn~(2+)/Cd~(2+)的积累有一定的作用;东南景天对Zn~(2+)和Cd~(2+)的吸收和积累存在不同的调控机制。
3.为方便比较锌超积累生态型东南景天在不同条件下的基因表达差异,需要克隆一个稳定表达的基因作为对照。在本试验中,利用设计的兼并引物通过PCR方法获得了一段actin基因的序列,然后根据获得的序列再通过3′-RACE和5′-RACE方法,得到了全长的actin基因序列;克隆得到的actin基因全长为1418bp,预测的编码区域为1131bp,编码377个氨基酸。
4利用cDNA-AFLP(cDNA扩增长度片段多态性:cDNA-amplified fragment lengthpolymorphism)方法,研究了我国原生的东南景开在锌超积累过程中的基因表达差异。结果表明:随着处理时间的增加,地上部锌含量也呈逐渐增加趋势;在5个时间点的多态性电泳图中,共出现4000多个条带,其中约有100个左右的转录差异片段(TDFs:transcript-derived fragments),并成功克隆了87个TDFs。;经数据库对比分析,其中有47个与已知或假定的基因存着明显的同源性,6个与未知功能的基因有同源性,34个与数据库中现有的基因没有明显的相似性。随机地从87个克隆片段中的选取23个做Real-time PCR验证,有16个表达谱与cDNA-AFLP完全相同;在38个与现有已知功能基因相似的片段中,按照功能可分成转录因子、胁迫响应、转录促进、细胞和组织代谢、光合作用、表达调节以及混合功能等几类;34个没有同源性的和15个与未知功能基因(假定的基因和未知功能的基因)同源的加在一起共有49个,占总数87个的56.3%。说明植物的超积累能力,与许多目前尚不清楚的基因有关。在38个能通过BLAST方法确定具体功能的TDFs中,包含有信号转导、胁迫响应、转录因子、表达调节等各种类型,表明东南景天锌超积累过程需要各种类型的酶系参与,锌超积累性状的形成包括复杂的信号通路和生理生化代谢途径。
Phytoremediation is defined as the use of plants to decontaminate and/or remove pollutants from the environment.It has emerged as an alternative technique for removing toxic metals from soil and offers the benefits of being in situ,cost effective and environmentally sustainable plants with high metal uptake and accumulation capacity have been termed as hyperaccumulator species.Sedum Alfredii Hance(S.Alfredii)was found in an old Pb/Zn mined area,Southern China and was originally identified as a new Zn/Cd co-hyperaccumulating plant,it was probably mutants from the non-hyperaccumulating ecotypes of S.Alfredii after a considerably long period(more than one thousand years)of inhabitation the highly heavy metal polluted environments.It has an extraordinary ability to tolerate and accumulate high concentration of environmental Zn and Cd,and with the characteristics of large biomass,fast growth speed,asexual propagation and perennial. Therefore,it is an ideal plant for studying mechanisms which are responsible for hyperaccumulation and phytoremediation practice.In the present study,we focused on generating dendrogram by using RAPD methods,and the analysis of the band sequences that only observed in HE as compared with other species of Sedum;otherwise, cDNA-amplified fragment length polymorphism(cDNA-AFLP)analysis was employed to identify the genes of Sedum alfredii H.that exhibit a modulated expression under Zn treatment applied in hydroponics culture;in addition,Zn and Cd concentrations in plants, reduced glutathione and Hydrogen Peroxide,total Sulphur were detected,and the role were analyzed in hyperaccumulations.The major results were summarized as follows:
1.Sedum alfredii Hance has been identified as a new Zn/Cd co-hyperaccumulating plant species from heavy metal polluted area native to China.However,little information is available for the genetic differences among this ecotype and its relatives.In this paper, genetic variation and relationship among several species of Sedum including hyperaccumulating ecotype(HE)and the non-hyperaccumulating ecotype(NHE)of S. alfredii H,well as Sedum including Sedum Makinoi Maxim(SMM),Sedum emarginatum Migo(SEM)、Sedum Sarmentoum Bunge(SSB),and Sedum Bulbiferum Makino:(SBM),was determined by random amplified polymorphic DNA(RAPD) analysis.Phytogenetic analysis suggested that they can be divided into two groups and the non-hyperaccumulating ecotype(NHE)of Sedum alfredii H is the nearest relatives of the hyperaccumulating ecotype(HE).However,genetics variation of HE and NHE is notable,which indicated that long term pollution of soil exerted great influenceon the genetic diversity and plant evolution.
2.Hyperaccumulation of heavy metals in plants is closely related to metal detoxification in plant cells to metal toxicity,however the mechanisms is not fully understood yet.In this study,the possible roles of superoxide(H_2O_2)in zinc and cadmium hyperaccumulation of Sedum alfredii Hance were analyzed.The results showed that Zn/Cd concentration in the shoots of Sedum afredii Hance increased linearly within 15 days,after then leveled off.Concentration of total GSH,total S,as well as H_2O_2 in the shoot also increased within 15 days correspondingly,and then decreased with further increasing time of metal treatment.Shoot total S and glutathione(GSH)were higher for Cd than for Zn treatment.However,reverse trends in shoot H_2O_2 concentration were noted,i.e.much higher H_1O_2 concentration was observed in Zn-treated plant shoot than in Cd-treated plant shoot.Increasing external S supply could increase Zn and Cd accumulation in Sedum alfredii H.These results suggested that S containing compounds coundn't be the main chelator of Zn or Cd in the hyperaccumulating ecotype of Sedum alfredii Hance.GSH may play an important role in the detoxification of dissociated Zn and Cd,and H_2O_2 may be a kind of signal molecule in correlation with Zn and/or Cd hyperaccumulation.
3.A pair of degenerate primers were designed using the plant actin sequences already available,actin genes from the hyperaccumulator Sedum alfredii H.were PCR-amplified, cloned,and sequenced.Then,both 5'-and 3'-rapid amplification of cDNA ends(RACE) were performed.The actin gene of full-length is 1418bp,and putative reading frame from 19bp to 1149bp(together 113 lbp).
4.cDNA-amplified fragment length polymorphism(cDNA-AFLP)analysis was employed to identify genes that exhibited an indusibleexpression under Zn treatment in Sedum alfredii H.grown in hydroponic culture,plants were treated for 3h,8h,24h and 1 weeks with 500μM Zn and untreated plants were used as controls.The results showed that there existed over 4000 expression bands appeared at different treatment times.Among over 100 transcript-derived fragments(TDFs)87 TDPs cloned as Zn responsive.. According to the analysis with genomic database,about were identified 47 TDFswere identified t are obviously homologous to the genes that have known or putative function, 6 TDFs were homologous to uncharacterized genes while 34 TDFs did not show significant matches to any genes in the Genodatabase.Real-time PCR analysis showed that the expression profiles of 16 in 23 TDFs which were randomly selected from 87 TDFs had high similarity to those with cDNA-AFLP.Thirty eight genes of known function TDFs coud be devided into different functional groups including transcriptional factors,expression regulators,and stress responding and transport facilitation genes,as well as genes involved in cellular metabolism and organization and the photosynthetic process.These results,suggest that a multitude of processes are implicated in Zn hyperaccumulating process in Sedum alfredii H..There have been 49 responsive genes identified in the HE of Sedum alfredii H composing of,34 known genes with blast and 15 unknown or putative genes,which accounted for 56.3%of total 87 genes,. implying Zn hyperaccumulation is associated with many related to the functions of many unkonown genes.According to BLAST analysis of the functions for the known 38 genes, zinc hyper-accumulation seemed to be controlled by different pathways including signal transduction,stress response,transcriptal factors,and expression regulation.These results indicate that the process of Zn hyperaccumulation could be regulated by a complex system of signal induction and physi-biochemical metablism.
1.植物长期生长在重金属污染的生境中,逐渐进化成不同的生态型,利用我国原生的Zn/Cd超积累生态型东南景天(Hyperaccumulated ecotype:HE)为材料,以与其亲缘关系较近的非超积累生态型东南景天(NHE)、圆叶景天(Sedum MakinoiMaxim:SMM)、凹叶景天(Sedum emarginatum Migo:SEM)、垂盆草(SedumSarmentoum Bunge:SSB)以及株芽景天(Sedum Bulbiferum Makino:SBM)为对照,利用RAPD方法,研究超积累生态型东南景天基因组变异的程度,并对部分超积累景天特有的PCR条带进行测序,分析特异基因片段与超积累的关系。遗传进化树分析表明:六种不同生态型的景天可以分成两组,其中两种东南景天为一组,另外四种为一组;超积累生态型和野生型东南景天有着相似的遗传背景,相似系数显示变异程度已超过种的界限。说明污染土壤对植物的进化有着非常大的影响;部分超积累景天特异PCR片段测序表明,ATP-硫酰化酶(ATP sulfuryase)、脱水调节相关蛋白及一些未知基因片段可能与超积累性状有一定的关系。
2.植物对重金属锌/镉超积累与过氧化氢产生关系的研究表明,在1000uM Zn~(2+)或200uM Cd~(2+)处理条件下,超积累生态型东南景天地上部Zn~(2+)的积累量高于Cd~(2+)的积累量。地上部Zn~(2+)/Cd~(2+)积累时间动力学显示在前15天,呈线形增加,随后积累速度下降。同样,地上部总硫、GSH以及H_2O_2的含量也是在前15天迅速增加,后10天逐渐下降。总硫和GSH的含量都是Cd~(2+)处理下较Zn~(2+)处理的高,而H_2O_2的含量则是Cd~(2+)处理下较Zn~(2+)处理的低;地上部Zn~(2+)含量变化与地上部GSH、总硫及H_2O_2变化趋势相似,而Cd~(2+)则是在第5天时含量就达到最大,以后开始逐渐下降。以上结果推测:含硫有机物可能不是超积累生态型东南景天Zn~(2+)/Cd~(2+)积累的主要配体,GSH的功能主要是解除体内游离态Zn~(2+)/Cd~(2+)的毒性,H_2O_2很可能是与超积累相关的一种信号分子。增加硫元素的供应对提高zn~(2+)/Cd~(2+)的积累有一定的作用;东南景天对Zn~(2+)和Cd~(2+)的吸收和积累存在不同的调控机制。
3.为方便比较锌超积累生态型东南景天在不同条件下的基因表达差异,需要克隆一个稳定表达的基因作为对照。在本试验中,利用设计的兼并引物通过PCR方法获得了一段actin基因的序列,然后根据获得的序列再通过3′-RACE和5′-RACE方法,得到了全长的actin基因序列;克隆得到的actin基因全长为1418bp,预测的编码区域为1131bp,编码377个氨基酸。
4利用cDNA-AFLP(cDNA扩增长度片段多态性:cDNA-amplified fragment lengthpolymorphism)方法,研究了我国原生的东南景开在锌超积累过程中的基因表达差异。结果表明:随着处理时间的增加,地上部锌含量也呈逐渐增加趋势;在5个时间点的多态性电泳图中,共出现4000多个条带,其中约有100个左右的转录差异片段(TDFs:transcript-derived fragments),并成功克隆了87个TDFs。;经数据库对比分析,其中有47个与已知或假定的基因存着明显的同源性,6个与未知功能的基因有同源性,34个与数据库中现有的基因没有明显的相似性。随机地从87个克隆片段中的选取23个做Real-time PCR验证,有16个表达谱与cDNA-AFLP完全相同;在38个与现有已知功能基因相似的片段中,按照功能可分成转录因子、胁迫响应、转录促进、细胞和组织代谢、光合作用、表达调节以及混合功能等几类;34个没有同源性的和15个与未知功能基因(假定的基因和未知功能的基因)同源的加在一起共有49个,占总数87个的56.3%。说明植物的超积累能力,与许多目前尚不清楚的基因有关。在38个能通过BLAST方法确定具体功能的TDFs中,包含有信号转导、胁迫响应、转录因子、表达调节等各种类型,表明东南景天锌超积累过程需要各种类型的酶系参与,锌超积累性状的形成包括复杂的信号通路和生理生化代谢途径。
Phytoremediation is defined as the use of plants to decontaminate and/or remove pollutants from the environment.It has emerged as an alternative technique for removing toxic metals from soil and offers the benefits of being in situ,cost effective and environmentally sustainable plants with high metal uptake and accumulation capacity have been termed as hyperaccumulator species.Sedum Alfredii Hance(S.Alfredii)was found in an old Pb/Zn mined area,Southern China and was originally identified as a new Zn/Cd co-hyperaccumulating plant,it was probably mutants from the non-hyperaccumulating ecotypes of S.Alfredii after a considerably long period(more than one thousand years)of inhabitation the highly heavy metal polluted environments.It has an extraordinary ability to tolerate and accumulate high concentration of environmental Zn and Cd,and with the characteristics of large biomass,fast growth speed,asexual propagation and perennial. Therefore,it is an ideal plant for studying mechanisms which are responsible for hyperaccumulation and phytoremediation practice.In the present study,we focused on generating dendrogram by using RAPD methods,and the analysis of the band sequences that only observed in HE as compared with other species of Sedum;otherwise, cDNA-amplified fragment length polymorphism(cDNA-AFLP)analysis was employed to identify the genes of Sedum alfredii H.that exhibit a modulated expression under Zn treatment applied in hydroponics culture;in addition,Zn and Cd concentrations in plants, reduced glutathione and Hydrogen Peroxide,total Sulphur were detected,and the role were analyzed in hyperaccumulations.The major results were summarized as follows:
1.Sedum alfredii Hance has been identified as a new Zn/Cd co-hyperaccumulating plant species from heavy metal polluted area native to China.However,little information is available for the genetic differences among this ecotype and its relatives.In this paper, genetic variation and relationship among several species of Sedum including hyperaccumulating ecotype(HE)and the non-hyperaccumulating ecotype(NHE)of S. alfredii H,well as Sedum including Sedum Makinoi Maxim(SMM),Sedum emarginatum Migo(SEM)、Sedum Sarmentoum Bunge(SSB),and Sedum Bulbiferum Makino:(SBM),was determined by random amplified polymorphic DNA(RAPD) analysis.Phytogenetic analysis suggested that they can be divided into two groups and the non-hyperaccumulating ecotype(NHE)of Sedum alfredii H is the nearest relatives of the hyperaccumulating ecotype(HE).However,genetics variation of HE and NHE is notable,which indicated that long term pollution of soil exerted great influenceon the genetic diversity and plant evolution.
2.Hyperaccumulation of heavy metals in plants is closely related to metal detoxification in plant cells to metal toxicity,however the mechanisms is not fully understood yet.In this study,the possible roles of superoxide(H_2O_2)in zinc and cadmium hyperaccumulation of Sedum alfredii Hance were analyzed.The results showed that Zn/Cd concentration in the shoots of Sedum afredii Hance increased linearly within 15 days,after then leveled off.Concentration of total GSH,total S,as well as H_2O_2 in the shoot also increased within 15 days correspondingly,and then decreased with further increasing time of metal treatment.Shoot total S and glutathione(GSH)were higher for Cd than for Zn treatment.However,reverse trends in shoot H_2O_2 concentration were noted,i.e.much higher H_1O_2 concentration was observed in Zn-treated plant shoot than in Cd-treated plant shoot.Increasing external S supply could increase Zn and Cd accumulation in Sedum alfredii H.These results suggested that S containing compounds coundn't be the main chelator of Zn or Cd in the hyperaccumulating ecotype of Sedum alfredii Hance.GSH may play an important role in the detoxification of dissociated Zn and Cd,and H_2O_2 may be a kind of signal molecule in correlation with Zn and/or Cd hyperaccumulation.
3.A pair of degenerate primers were designed using the plant actin sequences already available,actin genes from the hyperaccumulator Sedum alfredii H.were PCR-amplified, cloned,and sequenced.Then,both 5'-and 3'-rapid amplification of cDNA ends(RACE) were performed.The actin gene of full-length is 1418bp,and putative reading frame from 19bp to 1149bp(together 113 lbp).
4.cDNA-amplified fragment length polymorphism(cDNA-AFLP)analysis was employed to identify genes that exhibited an indusibleexpression under Zn treatment in Sedum alfredii H.grown in hydroponic culture,plants were treated for 3h,8h,24h and 1 weeks with 500μM Zn and untreated plants were used as controls.The results showed that there existed over 4000 expression bands appeared at different treatment times.Among over 100 transcript-derived fragments(TDFs)87 TDPs cloned as Zn responsive.. According to the analysis with genomic database,about were identified 47 TDFswere identified t are obviously homologous to the genes that have known or putative function, 6 TDFs were homologous to uncharacterized genes while 34 TDFs did not show significant matches to any genes in the Genodatabase.Real-time PCR analysis showed that the expression profiles of 16 in 23 TDFs which were randomly selected from 87 TDFs had high similarity to those with cDNA-AFLP.Thirty eight genes of known function TDFs coud be devided into different functional groups including transcriptional factors,expression regulators,and stress responding and transport facilitation genes,as well as genes involved in cellular metabolism and organization and the photosynthetic process.These results,suggest that a multitude of processes are implicated in Zn hyperaccumulating process in Sedum alfredii H..There have been 49 responsive genes identified in the HE of Sedum alfredii H composing of,34 known genes with blast and 15 unknown or putative genes,which accounted for 56.3%of total 87 genes,. implying Zn hyperaccumulation is associated with many related to the functions of many unkonown genes.According to BLAST analysis of the functions for the known 38 genes, zinc hyper-accumulation seemed to be controlled by different pathways including signal transduction,stress response,transcriptal factors,and expression regulation.These results indicate that the process of Zn hyperaccumulation could be regulated by a complex system of signal induction and physi-biochemical metablism.
引文
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