九孔鲍cDNA文库的构建、四种重要免疫相关蛋白的发现及其功能研究
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摘要
鲍是一类重要的经济贝类品种,但是近年来由于环境污染和病害的发生而导致鲍的产量大幅度下降。为了探索研究鲍免疫反应相关机制,为鲍病害防治提供理论基础,本研究构建了九孔鲍血淋巴细胞cDNA文库,首次筛选出了4个尚未报道的鲍免疫反应相关基因：组氨酸核苷结合蛋白、胞嘧啶核苷脱氨酶、泛素连接酶Rbx1和胸腺肽β。并利用现代生物信息学分析手段,分子生物学、免疫学及细胞生物学等技术对这4个鲍新基因的结构和功能进行了研究。
1.构建了九孔鲍血淋巴细胞cDNA文库
共得到310个EST序列,其中包含18个鲍新基因全长序列。通过生物信息学软件分析,这些基因分别涉及到鲍的细胞代谢和生长、转录调控、物质合成、免疫反应等。并把相关基因序列提交到GenBank数据库。
2.九孔鲍组氨酸核苷结合蛋白的鉴定与功能和机制的研究
九孔鲍组氨酸核苷结合蛋白(ab-HINT) cDNA全长为616 bp,氨基酸序列与哺乳动物HINT有较高的相似度,并且包含三个保守的活性位点区域。组织表达分布研究表明：ab-HINT在鲍不同组织中均有表达,表现了其功能的多样性,并且在鲍血淋巴细胞和鳃中的表达水平要明显高于其他组织,暗示了ab-HINT在鲍免疫反应中起到一定的作用。另外,LPS和Poly I：C的刺激均能诱导ab-HINT的表达水平上升,进一步说明了ab-HINT在鲍免疫反应中的作用。为了进一步研究ab-HINT的生物学活性及生理学作用,表达和纯化了ab-HINT蛋白,并利用质谱技术鉴定。分析型超速离心机分析结果指明该ab-HINT是一个同源二聚体。同时,高效液相色谱分析表明ab-HINT蛋白能够水解AMP-NH2。利用制备的抗体,用免疫组化和免疫荧光的方法,研究了ab-HINT在鲍组织细胞中的表达定位。在这些研究的基础上,我们进一步探索研究了ab-HINT对鲍血淋巴细胞及肿瘤细胞的凋亡作用及其作用机制。结果表明：ab-HINT能够诱导鲍血淋巴细胞凋亡,有可能是通过凋亡调控因子P53来实现的。另外,ab-HINT能够增加肿瘤细胞凋亡比例,该凋亡通路可能是通过上调P53的表达,同时伴随着抗凋亡蛋白Bcl-2的下调和促凋亡蛋白Bax的上调来实现的。
3.九孔鲍胞嘧啶核苷脱氨酶的鉴定及功能研究
九孔鲍胞嘧啶核苷脱氨酶(ab-CDA) cDNA全长为1752 bp,氨基酸序列中包含两个保守的活性位点区域和一个保守的半胱氨酸/组氨酸残基。序列比对及进化树分析表明ab-CDA属于同源四聚体家族。组织表达分布研究表明：ab-CDA在鲍不同组织中均有表达,并且在鲍血淋巴细胞中的表达水平显著高于其他组织。同时,LPS和Poly I:C的刺激均能诱导ab-CDA的表达水平上升,说明了ab-CDA在鲍免疫反应中的作用。为了进一步研究ab-CDA的生物学功能,表达和纯化了ab-CDA蛋白,并用质谱技术鉴定。分光光度计分析表明该蛋白具有胞嘧啶核苷脱氨酶活性,同时,利用免疫组化的方法,研究了ab-CDA在鲍组织细胞中的表达定位,结果表明：ab-CDA在胞质和胞核均有表达。
4.九孔鲍泛素连接酶Rbx1的鉴定及功能研究
九孔鲍泛素连接酶Rbx1 (ab-Rbxl)的cDNA全长为509 bp,氨基酸序列与其他Rbx1同源物有较高的相似度,并且ab-Rbx1氨基酸序列中含有Rbx1蛋白家族的特征区域ring box序列。组织表达分布研究表明：ab-Rbx1在鲍不同组织中均有表达,并且在鲍血淋巴细胞中的表达水平相对高于其他组织。同时,PHA的刺激能显著诱导ab-Rbx1的表达水平上升。说明了ab-Rbx1具有广泛的生物学功能,其中在免疫反应中起到一定的作用。为了进一步研究ab-Rbx1功能,表达和纯化了ab-Rbx1蛋白,并利用免疫组化和免疫荧光的方法,研究了ab-Rbx1在鲍组织细胞中的表达定位,结果表明ab-Rbx1在细胞质和细胞胞核均有一定表达,进一步说明了ab-Rbx1功能的多样性。另外,体外泛素化检测分析表明：ab-Rbx1具有泛素连接酶活性,并且这种活性依赖ATP,表明了泛素化修饰在软体动物鲍中的保守性。
5.九孔鲍胸腺肽β的鉴定与表达分析研究
九孔鲍胸腺肽β(ab-TMSB)的cDNA全长为499 bp,氨基酸序列与其他胸腺肽β同源物具有较高的相似性,并且序列中包含保守的肌动蛋白结合区,该区域是胸腺肽β蛋白家族的特征区。组织表达分布研究表明：ab-TMSB在鲍各检测的组织中均有一定的表达,表明ab-TMSB在鲍中具有广泛的生物学作用。并且ab-TMSB在血淋巴细胞中的表达要显著高于其他检测组织,暗示了ab-TMSB在鲍免疫反应中发挥一定的作用。进一步用实时定量PCR检测了ab-TMSB在大肠杆菌脂多糖(LPS)刺激后的表达变化,结果表明LPS能显著提高ab-TMSB的表达水平,初步揭示了ab-TMSB在鲍免疫反应中的作用。
Abalone is a type of important mollusc species for commercial production in the world. But in recent years, the global industry of abalone has been gradually decreased due to diseases and environmental pollution. Therefore, more investigation is required to aid in understanding the innate immune system of abalone for aquacμLture development. In this thesis, a cDNA library has been constructed and four novel immune-related genes have been selected in abalone (Haliotis diversicolor supertexta), including histidine triad nucleotide binding protein, cytidine deaminase, ubiquitin ligase RING boxl andβ-thymosin. Furthermore, functional analyses of these four genes have been carried out by bioinformatics methods, molecμLar biotechnology, immunology technology, cell biology and so on.
1. A cDNA library from abalone (H. diversicolor supertexta) was constructed
310 EST sequences were selected from the cDNA library including 18 novel genes. Based on bioinformatics analysis, these genes were annotated to be involved in different biological processes including cellular metabolic processes, cellular component organization and biosynthesis, signal transduction, immune defense and so on. The related genes sequences have been submitted to the GenBank.
2. Identification, function and mechanism research of ab-HINT
The full length cDNA of abalone (H. diversicolor supertexta) histidine triad nucleotide binding protein (ab-HINT) is 616 bp. It has higher similarity with mammalian HINT homologues and has three conserved domains. The results of the tissues distribution showed that the ab-HINT was ubiquitously expressed in abalone (H. diversicolor supertexta) tissues indicating that ab-HINT is involved in a wide variety of biological processes. Moreover, relatively higher amount of ab-HINT was observed in hemocyte and gills, which suggest that ab-HINT may be involved in the immune response of abalone (H. diversicolor supertexta). In addition, expression levels of ab-HINT increased quickly after LPS and Poly I:C challenge, which futher support our hypothesis. To further study the biological activity and physiological function of ab-HINT, the ab-HINT recombinant protein was expressed and purified, which was further identified by LC-MS/MS analyses. In addition, the structure of ab-HINT was studied by analytical ultracentrifμgation, which showed that ab-HINT is a homodimer. Analysis by HPLC demonstrates that the ab-HINT has the ability to hydrolyze AMP-NH2. To investigate the expression localization of ab-HINT in abalone tissues, we performed immunohistochemical staining and immunofluorescent staining with prepared anti-ab-HINT antibody. Based on the studies above, we examined the effects on cell apoptosis of ab-HINT. The results showed that ab-HINT can trigger hemocytes apoptosis and P53 is involved in this process. Moreover, ab-HINT can also induce human hepatocellular carcinoma (HCC) cells apoptosis via inducing an up-regulation of P53 expression coinciding with an up-regulation of the proapoptotic factor Bax and a concomitant down-regulation of the apoptosis inhibitor Bcl-2.
3. Identification and functional research of ab-CDA
The full length cDNA of abalone (H. diversicolor supertexta) cytidine deaminase (ab-CDA) is 1752 bp. It contains two conserved active site domains and conserved cysteine/histidine residues, which are the characteristics of CDA protein family. After sequence and phylogenetic tree analysis, we suggest that the ab-CDA is a homotetramer protein that belongs to the homotetrameric class of CDA superfamily. The results of the expression distribution revealed that ab-CDA was ubiquitously expressed in all examined tissues of abalone (H. diversicolor supertexta), which is consistent with the fact that ab-CDA is involved in a wide variety of biological processes. Moreover, the expression level in hemocyte is higher than that of other tissues, suggesting that ab-CDA may be involved in the immune response of abalone (H. diversicolor supertexta). The results that both LPS and Poly I:C can significantly up-regulate the expression level of ab-CDA further supported this point. To further study the biological activity and function of ab-CDA, the ab-CDA protein was expressed and purified, which was further confirmed by MS analyses. The enzymatic activity of ab-CDA studied by a spectrophotometric assay demonstrated that the ab-CDA fusion protein can deaminate cytidine effectively. Immunohistochemistry was performed to detect the intracellular localization of native ab-CDA in abalone (H. diversicolor supertexta) tissues. It was sμggested that ab-CDA is largely concentrated in the cytoplast and partially in the nuclei.
4. Identification and functional research of ab-Rbx1
The full length cDNA of abalone (H. diversicolor supertexta) ubiquitin ligase RING box1 (ab-Rbx1) is 509-bp. It has higher similarity with other Rbxl homologues and contains conserved cysteine/histidine residues. Study of the tissues distribution showed that ab-Rbx1 was ubiquitously expressed in abalone (H. diversicolor supertexta) tissues and a relatively high amount of ab-Rbx1 transcript was observed in hemocyte. Moreover, the expression levels of ab-Rbxl in hemocyte were increased quickly after PHA challenge. These results indicate that the ab-Rbx1 may be involved in a variety biological process in abalone (H. diversicolor supertexta) and may play a role in abalone (H. diversicolor supertexta) immune response. To further study the function of ab-Rbx1, the ab-Rbx1 was expressed snd purified. Immunohistochemical and immunofluorescent staining were performed with prepared antibody to investigate the expression localization of ab-Rbxl in abalone (H. diversicolor supertexta) tissues. The ab-Rbx1 was expressed predominantly in epithelial cells and localized both in the cytoplasmic and nuclear compartment. In addition, the activity of ab-Rbx1 was examined and the results showed that ab-Rbx1 had ubiquitin ligase activity, which was dependent on ATP.
5. Identification and expression analysis of ab-TMSB
The full length cDNA of abalone (H. diversicolor supertexta)β-thymosin (ab-TMSB) is 499 bp. It showed higher similarity with other TMSB homologous and contains the conserved actin-binding motif which is the characteristic ofβ-thymosin protein family. The tissue distribution profile revealed that the ab-TMSB was ubiquitously expressed in all examined tissues of abalone (H. diversicolor supertexta), which suggests that the ab-TMSB may be involved in a wide variety of biological processes. Moreover, a relatively higher expression level was observed in hemocyte, indicating that ab-TMSB may be involved in the immune response of abalone (H. diversicolor supertexta). In addition, expression levels of ab-TMSB were significantly up-regulated after LPS challenge, which futher suggest that ab-TMSB was involved in ablone (H. diversicolor supertexta) immune response.
1.构建了九孔鲍血淋巴细胞cDNA文库
共得到310个EST序列,其中包含18个鲍新基因全长序列。通过生物信息学软件分析,这些基因分别涉及到鲍的细胞代谢和生长、转录调控、物质合成、免疫反应等。并把相关基因序列提交到GenBank数据库。
2.九孔鲍组氨酸核苷结合蛋白的鉴定与功能和机制的研究
九孔鲍组氨酸核苷结合蛋白(ab-HINT) cDNA全长为616 bp,氨基酸序列与哺乳动物HINT有较高的相似度,并且包含三个保守的活性位点区域。组织表达分布研究表明：ab-HINT在鲍不同组织中均有表达,表现了其功能的多样性,并且在鲍血淋巴细胞和鳃中的表达水平要明显高于其他组织,暗示了ab-HINT在鲍免疫反应中起到一定的作用。另外,LPS和Poly I：C的刺激均能诱导ab-HINT的表达水平上升,进一步说明了ab-HINT在鲍免疫反应中的作用。为了进一步研究ab-HINT的生物学活性及生理学作用,表达和纯化了ab-HINT蛋白,并利用质谱技术鉴定。分析型超速离心机分析结果指明该ab-HINT是一个同源二聚体。同时,高效液相色谱分析表明ab-HINT蛋白能够水解AMP-NH2。利用制备的抗体,用免疫组化和免疫荧光的方法,研究了ab-HINT在鲍组织细胞中的表达定位。在这些研究的基础上,我们进一步探索研究了ab-HINT对鲍血淋巴细胞及肿瘤细胞的凋亡作用及其作用机制。结果表明：ab-HINT能够诱导鲍血淋巴细胞凋亡,有可能是通过凋亡调控因子P53来实现的。另外,ab-HINT能够增加肿瘤细胞凋亡比例,该凋亡通路可能是通过上调P53的表达,同时伴随着抗凋亡蛋白Bcl-2的下调和促凋亡蛋白Bax的上调来实现的。
3.九孔鲍胞嘧啶核苷脱氨酶的鉴定及功能研究
九孔鲍胞嘧啶核苷脱氨酶(ab-CDA) cDNA全长为1752 bp,氨基酸序列中包含两个保守的活性位点区域和一个保守的半胱氨酸/组氨酸残基。序列比对及进化树分析表明ab-CDA属于同源四聚体家族。组织表达分布研究表明：ab-CDA在鲍不同组织中均有表达,并且在鲍血淋巴细胞中的表达水平显著高于其他组织。同时,LPS和Poly I:C的刺激均能诱导ab-CDA的表达水平上升,说明了ab-CDA在鲍免疫反应中的作用。为了进一步研究ab-CDA的生物学功能,表达和纯化了ab-CDA蛋白,并用质谱技术鉴定。分光光度计分析表明该蛋白具有胞嘧啶核苷脱氨酶活性,同时,利用免疫组化的方法,研究了ab-CDA在鲍组织细胞中的表达定位,结果表明：ab-CDA在胞质和胞核均有表达。
4.九孔鲍泛素连接酶Rbx1的鉴定及功能研究
九孔鲍泛素连接酶Rbx1 (ab-Rbxl)的cDNA全长为509 bp,氨基酸序列与其他Rbx1同源物有较高的相似度,并且ab-Rbx1氨基酸序列中含有Rbx1蛋白家族的特征区域ring box序列。组织表达分布研究表明：ab-Rbx1在鲍不同组织中均有表达,并且在鲍血淋巴细胞中的表达水平相对高于其他组织。同时,PHA的刺激能显著诱导ab-Rbx1的表达水平上升。说明了ab-Rbx1具有广泛的生物学功能,其中在免疫反应中起到一定的作用。为了进一步研究ab-Rbx1功能,表达和纯化了ab-Rbx1蛋白,并利用免疫组化和免疫荧光的方法,研究了ab-Rbx1在鲍组织细胞中的表达定位,结果表明ab-Rbx1在细胞质和细胞胞核均有一定表达,进一步说明了ab-Rbx1功能的多样性。另外,体外泛素化检测分析表明：ab-Rbx1具有泛素连接酶活性,并且这种活性依赖ATP,表明了泛素化修饰在软体动物鲍中的保守性。
5.九孔鲍胸腺肽β的鉴定与表达分析研究
九孔鲍胸腺肽β(ab-TMSB)的cDNA全长为499 bp,氨基酸序列与其他胸腺肽β同源物具有较高的相似性,并且序列中包含保守的肌动蛋白结合区,该区域是胸腺肽β蛋白家族的特征区。组织表达分布研究表明：ab-TMSB在鲍各检测的组织中均有一定的表达,表明ab-TMSB在鲍中具有广泛的生物学作用。并且ab-TMSB在血淋巴细胞中的表达要显著高于其他检测组织,暗示了ab-TMSB在鲍免疫反应中发挥一定的作用。进一步用实时定量PCR检测了ab-TMSB在大肠杆菌脂多糖(LPS)刺激后的表达变化,结果表明LPS能显著提高ab-TMSB的表达水平,初步揭示了ab-TMSB在鲍免疫反应中的作用。
Abalone is a type of important mollusc species for commercial production in the world. But in recent years, the global industry of abalone has been gradually decreased due to diseases and environmental pollution. Therefore, more investigation is required to aid in understanding the innate immune system of abalone for aquacμLture development. In this thesis, a cDNA library has been constructed and four novel immune-related genes have been selected in abalone (Haliotis diversicolor supertexta), including histidine triad nucleotide binding protein, cytidine deaminase, ubiquitin ligase RING boxl andβ-thymosin. Furthermore, functional analyses of these four genes have been carried out by bioinformatics methods, molecμLar biotechnology, immunology technology, cell biology and so on.
1. A cDNA library from abalone (H. diversicolor supertexta) was constructed
310 EST sequences were selected from the cDNA library including 18 novel genes. Based on bioinformatics analysis, these genes were annotated to be involved in different biological processes including cellular metabolic processes, cellular component organization and biosynthesis, signal transduction, immune defense and so on. The related genes sequences have been submitted to the GenBank.
2. Identification, function and mechanism research of ab-HINT
The full length cDNA of abalone (H. diversicolor supertexta) histidine triad nucleotide binding protein (ab-HINT) is 616 bp. It has higher similarity with mammalian HINT homologues and has three conserved domains. The results of the tissues distribution showed that the ab-HINT was ubiquitously expressed in abalone (H. diversicolor supertexta) tissues indicating that ab-HINT is involved in a wide variety of biological processes. Moreover, relatively higher amount of ab-HINT was observed in hemocyte and gills, which suggest that ab-HINT may be involved in the immune response of abalone (H. diversicolor supertexta). In addition, expression levels of ab-HINT increased quickly after LPS and Poly I:C challenge, which futher support our hypothesis. To further study the biological activity and physiological function of ab-HINT, the ab-HINT recombinant protein was expressed and purified, which was further identified by LC-MS/MS analyses. In addition, the structure of ab-HINT was studied by analytical ultracentrifμgation, which showed that ab-HINT is a homodimer. Analysis by HPLC demonstrates that the ab-HINT has the ability to hydrolyze AMP-NH2. To investigate the expression localization of ab-HINT in abalone tissues, we performed immunohistochemical staining and immunofluorescent staining with prepared anti-ab-HINT antibody. Based on the studies above, we examined the effects on cell apoptosis of ab-HINT. The results showed that ab-HINT can trigger hemocytes apoptosis and P53 is involved in this process. Moreover, ab-HINT can also induce human hepatocellular carcinoma (HCC) cells apoptosis via inducing an up-regulation of P53 expression coinciding with an up-regulation of the proapoptotic factor Bax and a concomitant down-regulation of the apoptosis inhibitor Bcl-2.
3. Identification and functional research of ab-CDA
The full length cDNA of abalone (H. diversicolor supertexta) cytidine deaminase (ab-CDA) is 1752 bp. It contains two conserved active site domains and conserved cysteine/histidine residues, which are the characteristics of CDA protein family. After sequence and phylogenetic tree analysis, we suggest that the ab-CDA is a homotetramer protein that belongs to the homotetrameric class of CDA superfamily. The results of the expression distribution revealed that ab-CDA was ubiquitously expressed in all examined tissues of abalone (H. diversicolor supertexta), which is consistent with the fact that ab-CDA is involved in a wide variety of biological processes. Moreover, the expression level in hemocyte is higher than that of other tissues, suggesting that ab-CDA may be involved in the immune response of abalone (H. diversicolor supertexta). The results that both LPS and Poly I:C can significantly up-regulate the expression level of ab-CDA further supported this point. To further study the biological activity and function of ab-CDA, the ab-CDA protein was expressed and purified, which was further confirmed by MS analyses. The enzymatic activity of ab-CDA studied by a spectrophotometric assay demonstrated that the ab-CDA fusion protein can deaminate cytidine effectively. Immunohistochemistry was performed to detect the intracellular localization of native ab-CDA in abalone (H. diversicolor supertexta) tissues. It was sμggested that ab-CDA is largely concentrated in the cytoplast and partially in the nuclei.
4. Identification and functional research of ab-Rbx1
The full length cDNA of abalone (H. diversicolor supertexta) ubiquitin ligase RING box1 (ab-Rbx1) is 509-bp. It has higher similarity with other Rbxl homologues and contains conserved cysteine/histidine residues. Study of the tissues distribution showed that ab-Rbx1 was ubiquitously expressed in abalone (H. diversicolor supertexta) tissues and a relatively high amount of ab-Rbx1 transcript was observed in hemocyte. Moreover, the expression levels of ab-Rbxl in hemocyte were increased quickly after PHA challenge. These results indicate that the ab-Rbx1 may be involved in a variety biological process in abalone (H. diversicolor supertexta) and may play a role in abalone (H. diversicolor supertexta) immune response. To further study the function of ab-Rbx1, the ab-Rbx1 was expressed snd purified. Immunohistochemical and immunofluorescent staining were performed with prepared antibody to investigate the expression localization of ab-Rbxl in abalone (H. diversicolor supertexta) tissues. The ab-Rbx1 was expressed predominantly in epithelial cells and localized both in the cytoplasmic and nuclear compartment. In addition, the activity of ab-Rbx1 was examined and the results showed that ab-Rbx1 had ubiquitin ligase activity, which was dependent on ATP.
5. Identification and expression analysis of ab-TMSB
The full length cDNA of abalone (H. diversicolor supertexta)β-thymosin (ab-TMSB) is 499 bp. It showed higher similarity with other TMSB homologous and contains the conserved actin-binding motif which is the characteristic ofβ-thymosin protein family. The tissue distribution profile revealed that the ab-TMSB was ubiquitously expressed in all examined tissues of abalone (H. diversicolor supertexta), which suggests that the ab-TMSB may be involved in a wide variety of biological processes. Moreover, a relatively higher expression level was observed in hemocyte, indicating that ab-TMSB may be involved in the immune response of abalone (H. diversicolor supertexta). In addition, expression levels of ab-TMSB were significantly up-regulated after LPS challenge, which futher suggest that ab-TMSB was involved in ablone (H. diversicolor supertexta) immune response.
引文
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