Hg诱导和SA调节苜蓿氧化胁迫反应及miRNAs的分离、鉴定及表达分析
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摘要
汞已经成为农田有毒重金属污染的主要来源之一,它在植物中积累会破坏细胞功能,影响植物生长和发育。为了了解汞对植物的毒害作用,我们用1-40μMHgCl2或用20μM HgCl2连续处理紫花苜蓿,研究汞诱导氧化胁迫的机制。组织染色表明在汞胁迫下,紫花苜蓿根的分生区和伸长区均发生明显的膜脂过氧化作用,且膜的完整性受到破坏。根和叶中硫代巴比妥酸反应物质(TBARS)含量的变化证实了这一点。用活性聚丙烯酰胺凝胶电泳(PAGE)检测脂氧合酶,发现在紫花苜蓿根和叶中分别有两个和三个同工酶,在汞胁迫下变化不尽相同。随着汞浓度的上升,叶片中02-含量及叶和根中H202含量上升,NADH氧化酶的活性上升。为了研究汞胁迫下的生化反应,我们检测了紫花苜蓿中抗氧化酶如超氧化物歧化酶(SOD),过氧化物酶(POD),过氧化氢酶(CAT),抗坏血酸过氧化物酶(APX)和谷胱甘肽还原酶(GR)的活性。经汞处理后,紫花苜蓿叶中SOD, POD, APX, GR和CAT总活性上升,根中SOD和POD活性也是上升的。高浓度的汞(40μM)激活根中APX活性,而10-20μM HgCl2抑制了GR活性。活性PAGE分析表明在叶和根中分别检测到了5和3种SOD同工酶,7和10种POD同工酶,8和4种APX同工酶,但汞胁迫下变化不同。仅在叶片中检测到一种CAT带。我们也检测了抗氧化剂如抗坏血酸(ASC)和谷胱甘肽(GSH)的含量,结果表明汞胁迫下,ASC和GSH在根中含量下降,在叶片中积累。我们的研究表明汞胁迫下紫花苜蓿中02-和H202含量上升和抗氧化酶及抗氧化剂含量的变化相关,这些结果不但可以作为土壤受汞污染的生化指示剂,而且可以为研究植物耐汞机制提供重要的理论依据。
水杨酸(SA)作为信号分子介导许多生物和非生物胁迫诱导的生理反应。我们研究了SA在调节汞诱导的紫花苜蓿根部氧化胁迫中的作用。用0.2mM SA预处理12h再用10μM Hg2+处理24h后发现,SA能缓解汞对根的毒害,具体表现为降低了根的脂质过氧化产物,促进汞胁迫下根的生长。组织染色进一步证明SA缓解了汞对膜完整性的破坏。SA预处理提高了汞胁迫下的NADH氧化酶,APX和POD的活性,但SOD的活性稍有降低。我们检测了ASC, GSH和脯氨酸含量,发现经SA预处理再用汞处理比仅用汞处理的紫花苜蓿根中积累了更多的ASC, GSH和脯氨酸。所有这些结果表明SA在保护植物抵御汞诱导的氧化胁迫中发挥着重要的作用。
microRNAs (miRNAs)是植物和动物中一类调控基因转录后表达的短链、非编码的小分子RNAs (small RNA, sRNA).迄今为止,人们通过生物信息学和实验方法分离和鉴定到了大量的miRNAs,并登录在miRNAs数据库中(http://www.sanger.ac.uk/)。我们用生物信息学的方法从蒺藜苜蓿(Medicago truncatula)的ESTs和GSS中预测到了新的miRNAs及它们的靶基因。将拟南芥,水稻及其它植物中已知的miRNAs与蒺藜苜蓿ESTs和GSS数据库进行比对的同时,设置一系列严格的筛选标准,分析候选miRNAs序列特征,包括二级结构、茎环长度和miRNAs的保守性,最终筛选出了38个miRNAs.去除已登录的12个,我们在蒺藜苜蓿中一共发现了26个新的miRNAs,它们共代表15个家族。进一步将新鉴定的miRNAs和蒺藜苜蓿mRNA数据库进行比对,预测到了16个靶基因。分析表明上述大多数靶基因编码的产物为转录因子及重要代谢酶类,控制着植物生长发育,信号转导及各种胁迫反应。为了验证预测的正确性,我们用RT-PCR的方法,在蒺藜苜蓿根、茎、叶和花中验证到8个miRNAs,并分析了这些miRNAs在受汞胁迫的叶片中的表达谱。
尽管用生物信息学的方法可以分离和鉴定出很多miRNAs,但是其它非保守的miRNAs只有通过直接克隆法才能分离得到。为了获得新的响应胁迫的miRNAs,我们构建了一个经20μM HgCl2处理的蒺藜苜蓿sRNA文库,结果在蒺藜苜蓿中一共克隆到10个新的miRNAs,其中5个为植物中保守序列(1个在上述的生物信息学方法已经预测到),另外5个miRNAs在其它物种中还没有发现,这些miRNAs的生理功能有待于做进一步的鉴定。
Mercury has become one of the major sources of toxic heavy metal pollution in agricultural lands. Accumulation of mercury by plants may disrupt many cellular functions. To assess mercury toxicity, we performed the experiment focusing on the responses of alfalfa(Medicago sativa) to Hg+-induced oxidative stress. Alfalfa was treated with mercuric ions in a concentration-and time-dependent manner, and lipid peroxidation was studied biochemically as well as histochemically along with other physiological responses. Histochemical staining with Schiff's reagent and Evans blue revealed the peroxidation of membrane lipids and loss of plasma membrane integrity in the Hg-treated root apex. The histochemical observations were supported by the quantitative determinations of thiobarbituric acid reactive substances (TB ARS) in the roots and leaves of alfalfa. Analysis of lipoxygenase (LOX) activity by non-denaturing polyacrylamide gel electrophoresis (PAGE) showed that there were two isoforms in the root and three isoforms in the leaves of alfalfa, but they showed different patterns under the Hg exposure. The external concentrations of Hg2+were positively correlated with the generation of O2-in leaves and H2O2in leaves and roots. Treatment with Hg2+increased the activities of NADH oxidase. To understand the biochemical responses under the Hg stress, activities of antioxidant enzymes, superoxide dismutase (SOD), peroxidase (POD), catalase (CAT), ascorbate peroxidase (APX) and glutathione reductase (GR) were assayed. The total activities of SOD, POD, APX, GR and CAT were generally enhanced in leaves after exposure to Hg ions. Activities SOD and POD increased in roots after Hg treatment. Activity of APX was stimulated at relatively high concentration of Hg2+(40μM). In contrast, GR activity was depressed at higher concentrations of Hg+(10-20μM). Analysis of antioxidant enzymes activities in alfalfa by non-denaturing PAGE revealed five and three SOD isoforms, seven and ten POD isoforms, eight and four APX isoforms in leaves and roots, respectively. But they showed different patterns under Hg stress. Only one band of CAT was visualized in leaves. We also measured several antioxidative metabolites such as ascorbate (ASC) and glutathione (GSH). Treatment of seedlings with10-40μM Hg2+decreased the reduced ASC and GSH amounts in the roots but increased in the leaves. These results indicate that the increased levels of O2and H2O2under the Hg stress were closely linked to the changes of the activities of antioxidant enzymes and the content of antioxidative metabolites. The data not only provide the important information for better understanding of the tolerant mechanisms, but as well can be used as a bio-indicator for soil contamination by Hg.
Salicylic acid (SA) is a signal molecule mediating many biotic and environmental stress-induced physiological responses in plants. We investigated the role of SA in regulating Hg-induced oxidative stress in the roots of alfalfa. Plants pretreated with0.2mM SA for12h and subsequently exposed to10μM Hg2+for24h displayed attenuated toxicity to the roots. The SA-promoted root elongation was correlated with decreased lipid peroxidation in root cells. The ameliorating effect of SA was confirmed by the histochemical staining for the detection of loss of membrane integrity in Hg-treated roots. We show that treatment with0.2mM SA increased the activity of NADH oxidase, APX and POD in the roots exposed Hg. However, a slightly decreased SOD activity was observed in SA+Hg-treated roots when compared to those of Hg treatment alone. We also measured accumulation of ASC, GSH and proline in the roots of alfalfa and found that roots treated with S A in the presence of Hg accumulated more ASC, GSH and proline than those treated with Hg alone.
microRNAs (miRNAs) are a novel class of short, endogenous non-coding small RNAs that regulate gene post-transcriptional expression in bot plants and animals. Thus far, a growing number of miRNAs have been isolated and characterized from a variety of plant species by bioinformatical and experimental methods and deposited in the major miRNAs database (http://www.sanger.ac.uk/). Here we present a bioinformatic approach for ESTs-and GSS-wide prediction of novel miRNAs as well as their targets in Medicago truncatula. We initiated the prediction by using previously known miRNA sequences from Arabidopsis, rice and other plant species to blast the databases of M. truncatula ESTs and GSS to search for the potential miRNAs. We used the enlarged data set to analyze parameters of the plant precursors including secondary structure, stem length and conservation of miRNAs. A total of38potential miRNAs were detected following a variety of filtering criteria. After removal of the12overlapping miRNAs deposited in miRNA Registry (Release9.1),26miRNAs representing15families are found to be new. Using the newly identified miRNA sequences, we were able to further blast the M. truncatula mRNA database and detected16potential targets of miRNAs in M. truncatula. It was found that many miRNA targeted genes were predicted to encode transcription factors that regulate cell growth and development, signaling, and metabolism. To validate the predicted miRNAs in M. truncatula, we performed a RT-PCR based assay of mature miRNA expression. Eight miRNAs were identified in roots, stems, leaves and flowers of M. truncatula. Expression of the eight miRNAs was also performed on samples from the leaves subjected to mercury.
Although a number of plant miRNAs have been isolated by computational approach, many of other miRNAs, particularly those in response to abiotic stresses and non-conserved miRNAs appear to be found only by cloning approach. To identify novel abiotic stress-regulated miRNAs, we constructed a library of sRNAs from M. truncatula that were exposed to20μM HgCl2. Sequencing of the clones and subsequent analysis revealed10new miRNAs representing7families except one miRNA previously reported in M. truncatula miRNAs. Four of these were conserved in plant species, one of which was detected by bioinformatic approach above. Five new miRNAs of these from the library have no sequence conservation in any other plant species. The regulatory function of these cloned new miRNAs in M. truncatula under stress will be idetified further.
水杨酸(SA)作为信号分子介导许多生物和非生物胁迫诱导的生理反应。我们研究了SA在调节汞诱导的紫花苜蓿根部氧化胁迫中的作用。用0.2mM SA预处理12h再用10μM Hg2+处理24h后发现,SA能缓解汞对根的毒害,具体表现为降低了根的脂质过氧化产物,促进汞胁迫下根的生长。组织染色进一步证明SA缓解了汞对膜完整性的破坏。SA预处理提高了汞胁迫下的NADH氧化酶,APX和POD的活性,但SOD的活性稍有降低。我们检测了ASC, GSH和脯氨酸含量,发现经SA预处理再用汞处理比仅用汞处理的紫花苜蓿根中积累了更多的ASC, GSH和脯氨酸。所有这些结果表明SA在保护植物抵御汞诱导的氧化胁迫中发挥着重要的作用。
microRNAs (miRNAs)是植物和动物中一类调控基因转录后表达的短链、非编码的小分子RNAs (small RNA, sRNA).迄今为止,人们通过生物信息学和实验方法分离和鉴定到了大量的miRNAs,并登录在miRNAs数据库中(http://www.sanger.ac.uk/)。我们用生物信息学的方法从蒺藜苜蓿(Medicago truncatula)的ESTs和GSS中预测到了新的miRNAs及它们的靶基因。将拟南芥,水稻及其它植物中已知的miRNAs与蒺藜苜蓿ESTs和GSS数据库进行比对的同时,设置一系列严格的筛选标准,分析候选miRNAs序列特征,包括二级结构、茎环长度和miRNAs的保守性,最终筛选出了38个miRNAs.去除已登录的12个,我们在蒺藜苜蓿中一共发现了26个新的miRNAs,它们共代表15个家族。进一步将新鉴定的miRNAs和蒺藜苜蓿mRNA数据库进行比对,预测到了16个靶基因。分析表明上述大多数靶基因编码的产物为转录因子及重要代谢酶类,控制着植物生长发育,信号转导及各种胁迫反应。为了验证预测的正确性,我们用RT-PCR的方法,在蒺藜苜蓿根、茎、叶和花中验证到8个miRNAs,并分析了这些miRNAs在受汞胁迫的叶片中的表达谱。
尽管用生物信息学的方法可以分离和鉴定出很多miRNAs,但是其它非保守的miRNAs只有通过直接克隆法才能分离得到。为了获得新的响应胁迫的miRNAs,我们构建了一个经20μM HgCl2处理的蒺藜苜蓿sRNA文库,结果在蒺藜苜蓿中一共克隆到10个新的miRNAs,其中5个为植物中保守序列(1个在上述的生物信息学方法已经预测到),另外5个miRNAs在其它物种中还没有发现,这些miRNAs的生理功能有待于做进一步的鉴定。
Mercury has become one of the major sources of toxic heavy metal pollution in agricultural lands. Accumulation of mercury by plants may disrupt many cellular functions. To assess mercury toxicity, we performed the experiment focusing on the responses of alfalfa(Medicago sativa) to Hg+-induced oxidative stress. Alfalfa was treated with mercuric ions in a concentration-and time-dependent manner, and lipid peroxidation was studied biochemically as well as histochemically along with other physiological responses. Histochemical staining with Schiff's reagent and Evans blue revealed the peroxidation of membrane lipids and loss of plasma membrane integrity in the Hg-treated root apex. The histochemical observations were supported by the quantitative determinations of thiobarbituric acid reactive substances (TB ARS) in the roots and leaves of alfalfa. Analysis of lipoxygenase (LOX) activity by non-denaturing polyacrylamide gel electrophoresis (PAGE) showed that there were two isoforms in the root and three isoforms in the leaves of alfalfa, but they showed different patterns under the Hg exposure. The external concentrations of Hg2+were positively correlated with the generation of O2-in leaves and H2O2in leaves and roots. Treatment with Hg2+increased the activities of NADH oxidase. To understand the biochemical responses under the Hg stress, activities of antioxidant enzymes, superoxide dismutase (SOD), peroxidase (POD), catalase (CAT), ascorbate peroxidase (APX) and glutathione reductase (GR) were assayed. The total activities of SOD, POD, APX, GR and CAT were generally enhanced in leaves after exposure to Hg ions. Activities SOD and POD increased in roots after Hg treatment. Activity of APX was stimulated at relatively high concentration of Hg2+(40μM). In contrast, GR activity was depressed at higher concentrations of Hg+(10-20μM). Analysis of antioxidant enzymes activities in alfalfa by non-denaturing PAGE revealed five and three SOD isoforms, seven and ten POD isoforms, eight and four APX isoforms in leaves and roots, respectively. But they showed different patterns under Hg stress. Only one band of CAT was visualized in leaves. We also measured several antioxidative metabolites such as ascorbate (ASC) and glutathione (GSH). Treatment of seedlings with10-40μM Hg2+decreased the reduced ASC and GSH amounts in the roots but increased in the leaves. These results indicate that the increased levels of O2and H2O2under the Hg stress were closely linked to the changes of the activities of antioxidant enzymes and the content of antioxidative metabolites. The data not only provide the important information for better understanding of the tolerant mechanisms, but as well can be used as a bio-indicator for soil contamination by Hg.
Salicylic acid (SA) is a signal molecule mediating many biotic and environmental stress-induced physiological responses in plants. We investigated the role of SA in regulating Hg-induced oxidative stress in the roots of alfalfa. Plants pretreated with0.2mM SA for12h and subsequently exposed to10μM Hg2+for24h displayed attenuated toxicity to the roots. The SA-promoted root elongation was correlated with decreased lipid peroxidation in root cells. The ameliorating effect of SA was confirmed by the histochemical staining for the detection of loss of membrane integrity in Hg-treated roots. We show that treatment with0.2mM SA increased the activity of NADH oxidase, APX and POD in the roots exposed Hg. However, a slightly decreased SOD activity was observed in SA+Hg-treated roots when compared to those of Hg treatment alone. We also measured accumulation of ASC, GSH and proline in the roots of alfalfa and found that roots treated with S A in the presence of Hg accumulated more ASC, GSH and proline than those treated with Hg alone.
microRNAs (miRNAs) are a novel class of short, endogenous non-coding small RNAs that regulate gene post-transcriptional expression in bot plants and animals. Thus far, a growing number of miRNAs have been isolated and characterized from a variety of plant species by bioinformatical and experimental methods and deposited in the major miRNAs database (http://www.sanger.ac.uk/). Here we present a bioinformatic approach for ESTs-and GSS-wide prediction of novel miRNAs as well as their targets in Medicago truncatula. We initiated the prediction by using previously known miRNA sequences from Arabidopsis, rice and other plant species to blast the databases of M. truncatula ESTs and GSS to search for the potential miRNAs. We used the enlarged data set to analyze parameters of the plant precursors including secondary structure, stem length and conservation of miRNAs. A total of38potential miRNAs were detected following a variety of filtering criteria. After removal of the12overlapping miRNAs deposited in miRNA Registry (Release9.1),26miRNAs representing15families are found to be new. Using the newly identified miRNA sequences, we were able to further blast the M. truncatula mRNA database and detected16potential targets of miRNAs in M. truncatula. It was found that many miRNA targeted genes were predicted to encode transcription factors that regulate cell growth and development, signaling, and metabolism. To validate the predicted miRNAs in M. truncatula, we performed a RT-PCR based assay of mature miRNA expression. Eight miRNAs were identified in roots, stems, leaves and flowers of M. truncatula. Expression of the eight miRNAs was also performed on samples from the leaves subjected to mercury.
Although a number of plant miRNAs have been isolated by computational approach, many of other miRNAs, particularly those in response to abiotic stresses and non-conserved miRNAs appear to be found only by cloning approach. To identify novel abiotic stress-regulated miRNAs, we constructed a library of sRNAs from M. truncatula that were exposed to20μM HgCl2. Sequencing of the clones and subsequent analysis revealed10new miRNAs representing7families except one miRNA previously reported in M. truncatula miRNAs. Four of these were conserved in plant species, one of which was detected by bioinformatic approach above. Five new miRNAs of these from the library have no sequence conservation in any other plant species. The regulatory function of these cloned new miRNAs in M. truncatula under stress will be idetified further.
引文
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