高山离子芥铁蛋白基因CbFer的克隆、表达及其抗氧化功能分析
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摘要
高山离子芥(Chorispora bungeana Fisch. & C. A. Mey)是一种稀有的高山冰缘植物，具有很强抗寒能力，它不具备特殊的形态结构，但在生理和分子水平上是研究植物适应冷冻环境机制的理想材料。本文以高山离子芥为实验材料，克隆到一个全新的铁蛋白基因(CbFer)，在不同温度下(4℃、0℃和-4℃)对其进行低温胁迫后，系统性的检测了AsA-GSH系统和CbFer基因对低温的响应表达，重点对诱导CbFer基因表达的信号途径进行了初步研究。结果认为高山离子芥作为高山冰缘植物所特有的抗寒机制主要表现在以下几个方面：
1、高山离子芥铁蛋白基因(CbFer)全长975bp，有一个编码260个氨基酸的开放阅读框(ORF)。对应编码的蛋白分子量为29.17 kDa，等电点是5.44。在氨基酸结构域分析中，揭示了高山离子芥铁蛋白有类似铁蛋白双铁结构域并且具有铁蛋白铁结合区域的特征。
2、CbFer cDNA编码的铁蛋白亚基含有一段定位于叶绿体蛋白的转运肽。RT—PCR分析表明这个基因是组织特异性表达的基因，只能在叶片中检测到它的表达。这表明CbFer基因编码的铁蛋白功能在叶片中发挥。
3、检测这个基因在冷冻胁迫(4℃、0℃和-4℃)下的表达情况，发现它的转录物在4℃逐渐积累，而在0℃和-4℃的积累反而直接和迅速。这表明CbFer基因的表达与温度密切相关、对温度反应敏感，所以CbFer基因的表达是高山离子芥传递抵御低温胁迫信号的一个重要保护机制。随着胁迫强度的增大，植物启动这个保护机制就越迅速。这恰恰就反映了高山离子芥具有适应温度剧烈变化的能力。
4、铁鳌合剂(Ferrozine)处理明显的抑制了CbFer基因的表达，表明铁是调控CbFer基因表达的一个重要限制因子。但值得注意的是经铁鳌合剂处理后再进行冷冻胁迫又可以重新诱导CbFer基因的表达。这说明在冷冻情况下诱导CbFer表达的信号除了铁这个因素以外还可能包含其他因素。
5、用外源信号处理发现：过量铁和H_2O_2正调控CbFer基因的表达；ABA负调控CbFer基因的表达；SNP处理不能引起CbFer基因表达的变化。铁鳌合剂处理后引起的CbFer基因的表达抑制不能被外源ABA、Fe、SNP处理所恢复。但在同样情况下，用H_2O_2处理在一定程度上诱导了CbFer的表达。这表明CbFer
Chorispora bungeana Fisch. & C.A. Mey (C. bungeana) is a rare alpine subnival plant species that is highly capable of resisting freezing environment. Lacking special morphological structure, it is an ideal plant in studying physiological and molecular mechanism in resisting freezing stress. Ferritin cDNA, CbFer, was cloned from C. bungeana. The responses of AsA-GSH system and CbFer to temperature change were examined during the course of different temperature (4℃、0℃ and -4℃) stresses, the signal transduction way of the expression of CbFer was also investigated in this paper. The special cold resistance ability of C. bungeana shows in the following aspects:
1. CbFer gene is 975 bp in length with an open reading frame (ORF) of 260 amino acids, corresponding to a protein of predicted molecular mass 29.17 kDa and an isoelectric point of 5.44. The deduced amino acid sequence possesses ferritin-like diiron domain and the ferritin iron-binding regions signature.
2. CbFer cDNA codes for a ferritin subunit plus a chloroplast-targeting transit peptide. RT-PCR analysis revealed that CbFer was a tissue-specfic gene, the expression could only be detected in leaves. These results suggest that the protein coded by CbFer plays function only in leaves.
3. The expression patterns of CbFer gene were investigated in relation to cold stress. The CbFer gene transcript progressively increased under 4℃ and increased much more distinctly and quickly under 0℃ and -4℃. This result suggests there is a close relationship between the expression of CbFer and cold stress, CbFer expression is sensitive to low temperature. CbFer plays an important protective mechanism in signal transduction to resist cold stress in C. bungeana. Lower temperature could trigger the expression more quickly exactly, this phenomenon reflects fitly that C. bungeana has the ability to adapt to acute temperature change in the growth environment.
4. The expression of CbFer was strongly inhibited by treatment with iron chelating agent Ferrozine. This result indicates that iron is a main factor in the regulating of CbFer gene expression. It was remarkable that we found cold stress could induce the expression of CbFer after Ferrozine treatment, which means there is another signal except iron induces the expression of the gene during cold stress.
5. Two tested exogenous signals, excessive iron and H_2O_2, up-regulated the expression of CbFer gene. However, the expression of the gene was down-regulated
1、高山离子芥铁蛋白基因(CbFer)全长975bp，有一个编码260个氨基酸的开放阅读框(ORF)。对应编码的蛋白分子量为29.17 kDa，等电点是5.44。在氨基酸结构域分析中，揭示了高山离子芥铁蛋白有类似铁蛋白双铁结构域并且具有铁蛋白铁结合区域的特征。
2、CbFer cDNA编码的铁蛋白亚基含有一段定位于叶绿体蛋白的转运肽。RT—PCR分析表明这个基因是组织特异性表达的基因，只能在叶片中检测到它的表达。这表明CbFer基因编码的铁蛋白功能在叶片中发挥。
3、检测这个基因在冷冻胁迫(4℃、0℃和-4℃)下的表达情况，发现它的转录物在4℃逐渐积累，而在0℃和-4℃的积累反而直接和迅速。这表明CbFer基因的表达与温度密切相关、对温度反应敏感，所以CbFer基因的表达是高山离子芥传递抵御低温胁迫信号的一个重要保护机制。随着胁迫强度的增大，植物启动这个保护机制就越迅速。这恰恰就反映了高山离子芥具有适应温度剧烈变化的能力。
4、铁鳌合剂(Ferrozine)处理明显的抑制了CbFer基因的表达，表明铁是调控CbFer基因表达的一个重要限制因子。但值得注意的是经铁鳌合剂处理后再进行冷冻胁迫又可以重新诱导CbFer基因的表达。这说明在冷冻情况下诱导CbFer表达的信号除了铁这个因素以外还可能包含其他因素。
5、用外源信号处理发现：过量铁和H_2O_2正调控CbFer基因的表达；ABA负调控CbFer基因的表达；SNP处理不能引起CbFer基因表达的变化。铁鳌合剂处理后引起的CbFer基因的表达抑制不能被外源ABA、Fe、SNP处理所恢复。但在同样情况下，用H_2O_2处理在一定程度上诱导了CbFer的表达。这表明CbFer
Chorispora bungeana Fisch. & C.A. Mey (C. bungeana) is a rare alpine subnival plant species that is highly capable of resisting freezing environment. Lacking special morphological structure, it is an ideal plant in studying physiological and molecular mechanism in resisting freezing stress. Ferritin cDNA, CbFer, was cloned from C. bungeana. The responses of AsA-GSH system and CbFer to temperature change were examined during the course of different temperature (4℃、0℃ and -4℃) stresses, the signal transduction way of the expression of CbFer was also investigated in this paper. The special cold resistance ability of C. bungeana shows in the following aspects:
1. CbFer gene is 975 bp in length with an open reading frame (ORF) of 260 amino acids, corresponding to a protein of predicted molecular mass 29.17 kDa and an isoelectric point of 5.44. The deduced amino acid sequence possesses ferritin-like diiron domain and the ferritin iron-binding regions signature.
2. CbFer cDNA codes for a ferritin subunit plus a chloroplast-targeting transit peptide. RT-PCR analysis revealed that CbFer was a tissue-specfic gene, the expression could only be detected in leaves. These results suggest that the protein coded by CbFer plays function only in leaves.
3. The expression patterns of CbFer gene were investigated in relation to cold stress. The CbFer gene transcript progressively increased under 4℃ and increased much more distinctly and quickly under 0℃ and -4℃. This result suggests there is a close relationship between the expression of CbFer and cold stress, CbFer expression is sensitive to low temperature. CbFer plays an important protective mechanism in signal transduction to resist cold stress in C. bungeana. Lower temperature could trigger the expression more quickly exactly, this phenomenon reflects fitly that C. bungeana has the ability to adapt to acute temperature change in the growth environment.
4. The expression of CbFer was strongly inhibited by treatment with iron chelating agent Ferrozine. This result indicates that iron is a main factor in the regulating of CbFer gene expression. It was remarkable that we found cold stress could induce the expression of CbFer after Ferrozine treatment, which means there is another signal except iron induces the expression of the gene during cold stress.
5. Two tested exogenous signals, excessive iron and H_2O_2, up-regulated the expression of CbFer gene. However, the expression of the gene was down-regulated
引文
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