家蚕解毒酶基因表达调控研究
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摘要
在自然界中，生物体不断地与外界环境进行物质交换，以满足自身的生长、发育和生存需要。在这些物质中，有些是为了满足生物体营养所需，对生物体是有益的；有些却对生物体的生命有危害。为了生存，生物体必须具备一定的生理、生化，甚至是行为适应能力。通过体内的解毒酶系统催化分解代谢产物或异源物，起到解毒作用，是生物体主要且常见的一种适应机制。昆虫的解毒酶是一类异质酶系，能够代谢大量的内源或外源底物，这些解毒酶系包括：细胞色素P450酶系、谷胱甘肽-S-转移酶系和酯酶等。
应用geNorm和NormFinder软件首先对内参基因进行标准化分析以选择相对较稳定的内参基因，是相对定量过程中比较关键的步骤，然而即使选择了相对稳定的内参基因，由内参不稳定带来的误差仍是不容忽视的。而Dual spike-in qPCR通过向样品中加入外参基因，对目标基因进行全程跟踪标定，使校正结果较为准确。但是，该方法需同时测定DNA和RNA，使得该方法的工作量提高一倍，增加了实验成本和时间。为了解决这一问题，我们设想以Dual spike-in qPCR首先对已标准化的内参基因进行定量，再以此内参基因对目的基因进行校正的新的实验思路，结果表明该方法在能得到较准确定量结果的同时，降低了实验的工作量和成本，为实时荧光定量PCR技术的进一步发展提供了新的参考。
为了从宏观上解析家蚕体内主要解毒酶基因的分布情况以及外源物质对解毒酶基因转录水平的影响，我们采用Dual spike-in qPCR方法，测定了家蚕正常情况以及分别添食蜕皮激素和芸香苷条件下各组织中主要解毒酶基因的转录水平。结果表明：中肠中参与蜕皮激素代谢的基因相对较少，BmGST基因主要参与了中肠对芸香苷的代谢。脂肪体中参与蜕皮激素代谢的基因主要有CYP9a20, BmGSTo1, BmGSTz2等；参与芸香苷代谢的基因较多，以BmGST基因为主。马氏管中参与蜕皮激素代谢的基因主要有CYP6ab4, BmGSTs2, BmCarE15等；参与芸香苷代谢的基因主要有BmGSTo1。
为进一步了解家蚕解毒酶基因的生理功能，本研究选择了三个主要解毒酶基因：CYP6ab4、BmGSTd1、BmCarE-10。对上述基因设计了3个不同位点的siRNA，通过转染BmN细胞的方法筛选出干扰效果最显著的siRNA片段。结果表明：CYP6ab4-928，BmGSTd1-405，BmCarE-10-519的干扰效果最为显著，分别使各自干扰基因的转录水平下降到对照的26.5%，39.6%和17.9%，可用于进一步的体内RNAi实验。对家蚕五龄幼虫注射有效干扰siRNA片段，通过实时荧光定量PCR测定并分析经RNAi后家蚕解毒酶基因在中肠、脂肪体和马氏管3种组织中转录水平的变化。结果表明：家蚕五龄幼虫通过注射siRNA，3种被测基因表达都能够被成功抑制，且注射后48h比24h效果更为明显。为了解家蚕解毒酶基因RNAi后虫体对辛硫磷的抗性差异，对家蚕5龄幼虫分别注射siRNA片段后，添食浸有辛硫磷的桑叶，调查统计了不同处理下的累积致死率。结果显示：分别注射三个解毒酶基因siRNA48h后添食辛硫磷导致的虫体死亡率显著增高，分别达到88%、84%和89%。
为研究家蚕主要解毒酶基因的表达调控机制，对三个主要解毒酶基因：CYP6ab4、BmGSTd1、BmCarE-10的启动子区域进行了功能分析，构建了含荧光素酶报告基因和启动子不同长度顺次缺失片段的重组质粒，与内参载体pRL-TK共转染BmN细胞，用双荧光素酶报告基因检测系统检测启动子活性。结果表明：蜕皮激素能够上调CYP6ab4和BmGSTd1基因启动子的活性，而芸香苷能够上调BmGSTd1和BmCarE-10基因启动子的活性。家蚕CYP6ab4基因启动子5'单侧-827~-722bp是该基因启动子的主要活性区域，该区域内存在的BR-C Z可能在基因表达调控过程中起重要作用；BmGSTd1基因启动子5'单侧-1385~-1299bp区域内存在的Kr可能参与了基因的表达调控；BmCarE-10基因的转录活性区域是启动子5'单侧-705~-625bp，该区域内存在的Sn可能参与了基因的表达调控。
为了利用rAcMNPV作为基因介导的载体的双荧光定量表达载体系统对家蚕解毒酶基因启动子区域的功能进行分析。我们构建一个FHNLuc-A3RL双荧光质粒，其中萤火虫荧光素酶（FLuc）基因由解毒酶基因启动子带动，另一个由家蚕actin3启动子控制的海肾荧光素酶（RLuc）作内参基因。通过Bac-to-Bac系统制备重组杆状病毒，将这些重组病毒分别注射感染五龄起蚕，测定荧光素酶活性，从而分析启动子活性。结果表明：无论是在正常状态下还是在外源物质诱导下，各基因启动子在马氏管和脂肪体组织中的活性均高于其它组织，再次表明这两个组织在外源物质代谢过程中的重要性。而且其活性区域与BmN细胞中的测定结果基本一致，也再次证明了各活性区域内可能存在与基因表达调控相关的作用元件。
本研究通过定量PCR从宏观上分析了家蚕解素酶转录水平的基本情况；并通过双荧光素酶报告基因系统分析了三个重要解毒酶基因的启动子序列，得到了其中重要的调控区域，以及相关的蛋白因子。为家蚕解毒酶基因表达调控的研究提供了重要的理论依据。
In nature, in order to meet its own growth, development and survival needs,organisms make material exchanges with the external environment all the time. Some ofthese substances, in order to meet the organism nutrition requirement, is beneficial to theorganism, and some is harmful to them. In order to survive, the organism must have somephysiological, biochemical, and even behavioral adaptability. Catalyzing and decomposingmetabolites or heterologous material through the body's detoxification enzyme systemsplay a major and common adaptation mechanism. Insect detoxification enzymes are a classof heterogeneous enzymes which can metabolize endogenous or exogenous substrates,these detoxification enzyme mainly include: cytochrome P450, glutathione-S-transferaseand carboxylesterase.
In order to obtain a stable reference gene, standardization of reference gene by usingthe geNorm and NormFinder Software analysis is the key step in the process of relativequantification. However, the error caused by the relative quantification have not beeneliminated even choose a relatively stable reference gene. By adding an external referencegene into the sample, the target gene expression is calibrated and tracked, more accuratecorrection results are obtained by using Dual spike-in qPCR. However, according to theprinciple of the method, the workload is increased, and to some extent the experimentalcost and time is increased too. To solve this problem, we envision a new experimentalstrategy that we can quantify a standardized reference gene firstly by using Dual spike-inqPCR, and then use this gene to correct the target gene expression. The result showed thatthis strategy not only ensured the accurate results, but also reduced the experimental costand time. So, it promoted the further development and innovation of quantitative PCR tosome extent.
In order to understand the distribution of main detoxification genes in silkworm andthe impact of exogenous material on the expression of detoxification genes, thetranscription levels of detoxification genes in normal silkworm and silkworm fed by ecdysone and rutin were measured by using Dual spike-in PCR. The results show that lessgenes are involved in ecdysone metabolism in the midgut, and BmGST genes are maininvolved in rutin metabolism. The main genes involved in ecdysone metabolism areCYP9a20, BmGSTo1, BmGSTz2in fat body; more genes are involved in rutin metabolismsuch as BmGST genes. The main genes involved in ecdysone me tabolism are CYP6ab4,BmGSTs2, BmCarE-15in malpighian tubules, and BmGSTo1is mainly involved in rutinmetabolism.
To further understand the physiological functions of the detoxification genes, wedesigned three different siRNA for each gene in the present study, and the siRNAfragments with the most significant effect are filted. The results show that theCYP6ab4-928，BmGSTd1-405，BmCarE-10-519are most effective, the transcription levelof each gene decreased to26.5%,39.6%and17.9%respectively, so they can be used forfurther in vivo RNAi experiments.
In order to understand the impact of RNAi on silkworm gene expression in vivo, theeffective siRNA were injected into fifth instar silkworm larvae, and then we measured andanalyzed the changes of transcription levels of detoxification genes in mid gut, fat bodyand malpighian tubule by using real-time PCR. Experimental results show that theexpression of three measured genes can successfully be suppressed, and the extents are notsame. The effect at48h after injection is more obvious than24h. We surveyed thecumulative mortality rate of silkworms after feeding phoxim, and the results show that themortality was significantly increased to88%,84%and89%, respectively at48h afterinjection.
For the study on the mechanism of expression and regulation of main detoxificationgenes, we analyzed the function of promoter region of CYP6ab4, BmGSTd1andBmCarE-10genes, and constructed recombinant plasmids containing the luciferasereporter gene and promoter. The internal control vector pRL-TK was co-transfected intoBmN cell with recombinant plasmids, promoter activity was detected by using dualluciferase reporter gene assay system. The results show that the promoter activity ofCYP6ab4and BmGSTd1genes were raised by ecdysone, and rutin can raise the promoteractivity of BmGSTd1and BmCarE-10genes. The promoter activity area of CYP6ab4,BmGSTd1and BmCarE-10gene exists in the respective promoter5' unilateral-827~-722bp,-1385~-1299bp and-705~-625bp of various regions respectively, and the presence of the BR-C Z, Kr and Sn may play an important role in the process of gene expressionregulation.
We use rAcMNPV as a carrier of the gene-mediated fluorescence quantitativeexpression vector system to analyze the functions of the detoxification gene promoterregion. Constract a dual fluorescent plasmid of FHNLuc-A3RL, in which Fluc gene isdriven by detoxification gene promoter, and Rluc is driven by A3promoter. We preparedrecombinant baculovious through the Bac-to-Bac system, and injected them into fifth instarsilkworm, and analysis the luciferase activity. The results show that the of activities ofthree gene promoters in malpighian tubule and fat body are higher than in other tissues,and the promoter activity area is consistent with the measured results in BmN cells.
This study analyzed the whole basic transcription situation of detoxification genes insilkworm by using real-time PCR, and analyzed the promoter sequence of three importantdetoxification genes by using dual luciferase reporter gene assay system. Importantregulatory regions and associated protein factors were obtained. This study providedimportant theoretical basis for the study of the regulation of gene expression in silkwormdetoxification enzymes.
应用geNorm和NormFinder软件首先对内参基因进行标准化分析以选择相对较稳定的内参基因，是相对定量过程中比较关键的步骤，然而即使选择了相对稳定的内参基因，由内参不稳定带来的误差仍是不容忽视的。而Dual spike-in qPCR通过向样品中加入外参基因，对目标基因进行全程跟踪标定，使校正结果较为准确。但是，该方法需同时测定DNA和RNA，使得该方法的工作量提高一倍，增加了实验成本和时间。为了解决这一问题，我们设想以Dual spike-in qPCR首先对已标准化的内参基因进行定量，再以此内参基因对目的基因进行校正的新的实验思路，结果表明该方法在能得到较准确定量结果的同时，降低了实验的工作量和成本，为实时荧光定量PCR技术的进一步发展提供了新的参考。
为了从宏观上解析家蚕体内主要解毒酶基因的分布情况以及外源物质对解毒酶基因转录水平的影响，我们采用Dual spike-in qPCR方法，测定了家蚕正常情况以及分别添食蜕皮激素和芸香苷条件下各组织中主要解毒酶基因的转录水平。结果表明：中肠中参与蜕皮激素代谢的基因相对较少，BmGST基因主要参与了中肠对芸香苷的代谢。脂肪体中参与蜕皮激素代谢的基因主要有CYP9a20, BmGSTo1, BmGSTz2等；参与芸香苷代谢的基因较多，以BmGST基因为主。马氏管中参与蜕皮激素代谢的基因主要有CYP6ab4, BmGSTs2, BmCarE15等；参与芸香苷代谢的基因主要有BmGSTo1。
为进一步了解家蚕解毒酶基因的生理功能，本研究选择了三个主要解毒酶基因：CYP6ab4、BmGSTd1、BmCarE-10。对上述基因设计了3个不同位点的siRNA，通过转染BmN细胞的方法筛选出干扰效果最显著的siRNA片段。结果表明：CYP6ab4-928，BmGSTd1-405，BmCarE-10-519的干扰效果最为显著，分别使各自干扰基因的转录水平下降到对照的26.5%，39.6%和17.9%，可用于进一步的体内RNAi实验。对家蚕五龄幼虫注射有效干扰siRNA片段，通过实时荧光定量PCR测定并分析经RNAi后家蚕解毒酶基因在中肠、脂肪体和马氏管3种组织中转录水平的变化。结果表明：家蚕五龄幼虫通过注射siRNA，3种被测基因表达都能够被成功抑制，且注射后48h比24h效果更为明显。为了解家蚕解毒酶基因RNAi后虫体对辛硫磷的抗性差异，对家蚕5龄幼虫分别注射siRNA片段后，添食浸有辛硫磷的桑叶，调查统计了不同处理下的累积致死率。结果显示：分别注射三个解毒酶基因siRNA48h后添食辛硫磷导致的虫体死亡率显著增高，分别达到88%、84%和89%。
为研究家蚕主要解毒酶基因的表达调控机制，对三个主要解毒酶基因：CYP6ab4、BmGSTd1、BmCarE-10的启动子区域进行了功能分析，构建了含荧光素酶报告基因和启动子不同长度顺次缺失片段的重组质粒，与内参载体pRL-TK共转染BmN细胞，用双荧光素酶报告基因检测系统检测启动子活性。结果表明：蜕皮激素能够上调CYP6ab4和BmGSTd1基因启动子的活性，而芸香苷能够上调BmGSTd1和BmCarE-10基因启动子的活性。家蚕CYP6ab4基因启动子5'单侧-827~-722bp是该基因启动子的主要活性区域，该区域内存在的BR-C Z可能在基因表达调控过程中起重要作用；BmGSTd1基因启动子5'单侧-1385~-1299bp区域内存在的Kr可能参与了基因的表达调控；BmCarE-10基因的转录活性区域是启动子5'单侧-705~-625bp，该区域内存在的Sn可能参与了基因的表达调控。
为了利用rAcMNPV作为基因介导的载体的双荧光定量表达载体系统对家蚕解毒酶基因启动子区域的功能进行分析。我们构建一个FHNLuc-A3RL双荧光质粒，其中萤火虫荧光素酶（FLuc）基因由解毒酶基因启动子带动，另一个由家蚕actin3启动子控制的海肾荧光素酶（RLuc）作内参基因。通过Bac-to-Bac系统制备重组杆状病毒，将这些重组病毒分别注射感染五龄起蚕，测定荧光素酶活性，从而分析启动子活性。结果表明：无论是在正常状态下还是在外源物质诱导下，各基因启动子在马氏管和脂肪体组织中的活性均高于其它组织，再次表明这两个组织在外源物质代谢过程中的重要性。而且其活性区域与BmN细胞中的测定结果基本一致，也再次证明了各活性区域内可能存在与基因表达调控相关的作用元件。
本研究通过定量PCR从宏观上分析了家蚕解素酶转录水平的基本情况；并通过双荧光素酶报告基因系统分析了三个重要解毒酶基因的启动子序列，得到了其中重要的调控区域，以及相关的蛋白因子。为家蚕解毒酶基因表达调控的研究提供了重要的理论依据。
In nature, in order to meet its own growth, development and survival needs,organisms make material exchanges with the external environment all the time. Some ofthese substances, in order to meet the organism nutrition requirement, is beneficial to theorganism, and some is harmful to them. In order to survive, the organism must have somephysiological, biochemical, and even behavioral adaptability. Catalyzing and decomposingmetabolites or heterologous material through the body's detoxification enzyme systemsplay a major and common adaptation mechanism. Insect detoxification enzymes are a classof heterogeneous enzymes which can metabolize endogenous or exogenous substrates,these detoxification enzyme mainly include: cytochrome P450, glutathione-S-transferaseand carboxylesterase.
In order to obtain a stable reference gene, standardization of reference gene by usingthe geNorm and NormFinder Software analysis is the key step in the process of relativequantification. However, the error caused by the relative quantification have not beeneliminated even choose a relatively stable reference gene. By adding an external referencegene into the sample, the target gene expression is calibrated and tracked, more accuratecorrection results are obtained by using Dual spike-in qPCR. However, according to theprinciple of the method, the workload is increased, and to some extent the experimentalcost and time is increased too. To solve this problem, we envision a new experimentalstrategy that we can quantify a standardized reference gene firstly by using Dual spike-inqPCR, and then use this gene to correct the target gene expression. The result showed thatthis strategy not only ensured the accurate results, but also reduced the experimental costand time. So, it promoted the further development and innovation of quantitative PCR tosome extent.
In order to understand the distribution of main detoxification genes in silkworm andthe impact of exogenous material on the expression of detoxification genes, thetranscription levels of detoxification genes in normal silkworm and silkworm fed by ecdysone and rutin were measured by using Dual spike-in PCR. The results show that lessgenes are involved in ecdysone metabolism in the midgut, and BmGST genes are maininvolved in rutin metabolism. The main genes involved in ecdysone metabolism areCYP9a20, BmGSTo1, BmGSTz2in fat body; more genes are involved in rutin metabolismsuch as BmGST genes. The main genes involved in ecdysone me tabolism are CYP6ab4,BmGSTs2, BmCarE-15in malpighian tubules, and BmGSTo1is mainly involved in rutinmetabolism.
To further understand the physiological functions of the detoxification genes, wedesigned three different siRNA for each gene in the present study, and the siRNAfragments with the most significant effect are filted. The results show that theCYP6ab4-928，BmGSTd1-405，BmCarE-10-519are most effective, the transcription levelof each gene decreased to26.5%,39.6%and17.9%respectively, so they can be used forfurther in vivo RNAi experiments.
In order to understand the impact of RNAi on silkworm gene expression in vivo, theeffective siRNA were injected into fifth instar silkworm larvae, and then we measured andanalyzed the changes of transcription levels of detoxification genes in mid gut, fat bodyand malpighian tubule by using real-time PCR. Experimental results show that theexpression of three measured genes can successfully be suppressed, and the extents are notsame. The effect at48h after injection is more obvious than24h. We surveyed thecumulative mortality rate of silkworms after feeding phoxim, and the results show that themortality was significantly increased to88%,84%and89%, respectively at48h afterinjection.
For the study on the mechanism of expression and regulation of main detoxificationgenes, we analyzed the function of promoter region of CYP6ab4, BmGSTd1andBmCarE-10genes, and constructed recombinant plasmids containing the luciferasereporter gene and promoter. The internal control vector pRL-TK was co-transfected intoBmN cell with recombinant plasmids, promoter activity was detected by using dualluciferase reporter gene assay system. The results show that the promoter activity ofCYP6ab4and BmGSTd1genes were raised by ecdysone, and rutin can raise the promoteractivity of BmGSTd1and BmCarE-10genes. The promoter activity area of CYP6ab4,BmGSTd1and BmCarE-10gene exists in the respective promoter5' unilateral-827~-722bp,-1385~-1299bp and-705~-625bp of various regions respectively, and the presence of the BR-C Z, Kr and Sn may play an important role in the process of gene expressionregulation.
We use rAcMNPV as a carrier of the gene-mediated fluorescence quantitativeexpression vector system to analyze the functions of the detoxification gene promoterregion. Constract a dual fluorescent plasmid of FHNLuc-A3RL, in which Fluc gene isdriven by detoxification gene promoter, and Rluc is driven by A3promoter. We preparedrecombinant baculovious through the Bac-to-Bac system, and injected them into fifth instarsilkworm, and analysis the luciferase activity. The results show that the of activities ofthree gene promoters in malpighian tubule and fat body are higher than in other tissues,and the promoter activity area is consistent with the measured results in BmN cells.
This study analyzed the whole basic transcription situation of detoxification genes insilkworm by using real-time PCR, and analyzed the promoter sequence of three importantdetoxification genes by using dual luciferase reporter gene assay system. Importantregulatory regions and associated protein factors were obtained. This study providedimportant theoretical basis for the study of the regulation of gene expression in silkwormdetoxification enzymes.
引文
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