中华鳖免疫球蛋白连接链和重链基因与抗体产生规律
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摘要
中华鳖(Pelodiscus sinensis)是我国重要的淡水养殖动物,对其免疫系统的研究不仅在脊椎动物发育与比较免疫学方面具有重要意义,而且在中华鳖病害防治方面也具有重要的价值。本论文对编码中华鳖免疫球蛋白连接链(Joining chain)和免疫球蛋白M、D、Y的重链cDNA进行克隆,并揭示这些抗体的表达规律。此外,对中华鳖IgM、IgD和IgY阳性细胞的发生进行了研究。
用RACE-PCR方法扩增出中华鳖免疫球蛋白J(Joining chain)链全长cDNA。其cDNA全长为2,347 nt,包含5′非编码区30 nt,3′非编码区1,834 nt,开放阅读框480 nt,编码160个氨基酸。序列比对分析表明中华鳖免疫球蛋白J链基因与海龟的相似性高达80.7%,与鸡、爪蟾、牛蛙和鳐的相似性分别是71.3%、57.6%、49.4%和35.8%。结合进化树的结果分析,表明中华鳖免疫球蛋白J链基因同鸡的亲缘关系高于同两栖类的关系。用RT-PCR方法,检测到J链基因在中华鳖胚胎受精后5天就有表达,而且持续增加。免疫印迹实验和免疫组化分析表明J链基因组成型表达于中华鳖免疫器官和组织中。荧光定量PCR显示灭活的Aeromonas hydrophila strain T4可诱导中华鳖免疫球蛋白J链mRNA在脾脏、肾脏和血液中有较高的上调表达,表明免疫球蛋白J链基因在中华鳖早期发育阶段和抵抗微生物感染免疫应答过程中具有重要作用。
到目前为止,只有三种免疫球蛋白分子IgM、IgD和IgA在爬行动物中报道。本研究中,我们克隆了中华鳖IgM、IgD和一种新的免疫球蛋白分子IgY,其中IgM和IgY同其它物种的相应的Ig分子具有相似的特征,而IgD分子结构不同于其它物种,由6个不重复的恒定区组成。利用RT-PCR检测到IgM和IgD基因在中华鳖胚胎受精后7天就有表达,而且表达量随着发育持续增加。但IgY在孵化后90天才在脾脏和肾脏中检测到微弱的表达。通过免疫印迹实验,在中华鳖主要免疫器官和组织中都检测到三种免疫球蛋白分子IgM、IgD和IgY的表达。免疫组化检测到中华鳖主要免疫器官,如脾脏、肾脏和肠中均有IgM、IgD和IgY阳性细胞。而且,实时荧光定量PCR和间接ELISA分析了灭活嗜水气单胞菌A.hydrophila诱导后中华鳖脾脏、肾脏和血液中各类Ig基因转录水平和血靖中IgM、IgD和IgY抗体表达水平的变化,结果显示均有较高的表达上调。利用流式细胞术分析发现只有IgM在脾脏、肾脏和外周血白细胞中有较高的阳性比例。原位杂交检测到IgM阳性细胞首先在孵化后18天出现在中华鳖脾脏中,随后在28天和64天的肾脏和肠中检测到,而IgD和IgY阳性细胞出现时间要晚于IgM。
Chinese soft-shelled turtle, Pelodiscus sinensis is one of important reptile species in aquaculture industry of China. The understanding of its immune system is not only important and necessary for developmental and comparative immunity in vertebrates, but also for aquacultural purpose. The present study aims to clone the cDNAs encoding the heavy chains and joining chain of IgM, IgD and IgY in Chinese soft-shelled turtle, and to analyze their expression at mRNA and protein levels. Furthermore, the ontogeny of IgM-, IgD- and IgY- producing cells in lymphoid tissues of the turtle was also analyzed. The cDNA sequence of J chain has been cloned from the Chinese soft-shelled turtle by reverse transcription (RT)-PCR and rapid amplification of cDNA ends (RACE). The cDNA sequence is 2,347 nt in length and contains an open reading frame of 480 nt encoding 160 aa including the signal sequence. The deduced amino acid sequence has a high degree of homology with that of an already reported turtle J chain (80.7%), and of chicken (71.3%), and a low degree of homology with those of amphibians (49.4% and 57.6%); but it is relatively low when compared with the sequence reported from skate (35.8%). It is thus implied that the J chain of P. sinensis is much more similar to avian, as also revealed in the phylogenetic tree. The expression of J chain was detected at 5 days past fertilization (dpf) by RT-PCR, and its expression increased following further developmental stages. The expression of J chain was detected in almost all organs examined at protein levels by Western blotting, except in fat tissue. The J chain-producing cells were detected in lymphoid-related tissues such as in spleen, kidney and intestine by using immunohistochemistry. By using real-time quantitative RT-PCR analysis, a significant up-regulation of J chain transcripts was observed in spleen, kidney and blood of the turtle injected with in-activated Aeromonas hydrophila T4, indicating the important immune role of J chain at early development and in response to bacterial infection.
Only three Ig isotypes, IgM, IgD, and IgA, were previously known in reptiles. Here, we describe IgM, IgD and a novel immunoglobulin heavy-chain isotypeυ(IgY) in Chinese soft-shelled turtle. The IgM and IgY genes of the turtle are characteristically similar to those described in other species, whereas IgD gene differs from IgD reported in other vertebrates. The turtle IgD contains 6 immunoglobulin heavy chain domains without evidence of intragenic duplications of exons as described in teleost IgD genes. The expression of IgM and IgD was detected early during embryonic development after fertilization, and their expression increased following further developmental stages. However, immunoglobulin Y could not be detected in spleen and kidney until 90 days after hatching out. Western blotting analysis demonstrated that the protein of IgM, IgD and IgY were expressed in almost all organs examined in adult turtles. The IgM-, IgD- and IgY- producing cells in lymphoid-related tissues such as in spleen, kidney and intestine, were investigated by using histochemistry. The transcriptional kinetics of the three Ig molecules in spleen, kidney and blood and antibody response kinetics in serum after bacterial pathogen stimulation were also analyzed by using real-time PCR and by using indirected enzyme-linked immunosorbent assay (ELISA), respectively. A significant up-regulation of J chain was observed at both mRNA and protein levels. Using flow cytometry, a higher proportion of IgM-pruducing cells was observed in spleen, peripheral blood and kidney when compared to proportions of IgD-and IgY- positive cells. The IgM-producing cells was firstly observed in spleen of P. sinensis at 18 dph, and then in kidney and intestine at 28 and 64 dph, respectively. However, IgD and IgY positive cells were detected much later.
用RACE-PCR方法扩增出中华鳖免疫球蛋白J(Joining chain)链全长cDNA。其cDNA全长为2,347 nt,包含5′非编码区30 nt,3′非编码区1,834 nt,开放阅读框480 nt,编码160个氨基酸。序列比对分析表明中华鳖免疫球蛋白J链基因与海龟的相似性高达80.7%,与鸡、爪蟾、牛蛙和鳐的相似性分别是71.3%、57.6%、49.4%和35.8%。结合进化树的结果分析,表明中华鳖免疫球蛋白J链基因同鸡的亲缘关系高于同两栖类的关系。用RT-PCR方法,检测到J链基因在中华鳖胚胎受精后5天就有表达,而且持续增加。免疫印迹实验和免疫组化分析表明J链基因组成型表达于中华鳖免疫器官和组织中。荧光定量PCR显示灭活的Aeromonas hydrophila strain T4可诱导中华鳖免疫球蛋白J链mRNA在脾脏、肾脏和血液中有较高的上调表达,表明免疫球蛋白J链基因在中华鳖早期发育阶段和抵抗微生物感染免疫应答过程中具有重要作用。
到目前为止,只有三种免疫球蛋白分子IgM、IgD和IgA在爬行动物中报道。本研究中,我们克隆了中华鳖IgM、IgD和一种新的免疫球蛋白分子IgY,其中IgM和IgY同其它物种的相应的Ig分子具有相似的特征,而IgD分子结构不同于其它物种,由6个不重复的恒定区组成。利用RT-PCR检测到IgM和IgD基因在中华鳖胚胎受精后7天就有表达,而且表达量随着发育持续增加。但IgY在孵化后90天才在脾脏和肾脏中检测到微弱的表达。通过免疫印迹实验,在中华鳖主要免疫器官和组织中都检测到三种免疫球蛋白分子IgM、IgD和IgY的表达。免疫组化检测到中华鳖主要免疫器官,如脾脏、肾脏和肠中均有IgM、IgD和IgY阳性细胞。而且,实时荧光定量PCR和间接ELISA分析了灭活嗜水气单胞菌A.hydrophila诱导后中华鳖脾脏、肾脏和血液中各类Ig基因转录水平和血靖中IgM、IgD和IgY抗体表达水平的变化,结果显示均有较高的表达上调。利用流式细胞术分析发现只有IgM在脾脏、肾脏和外周血白细胞中有较高的阳性比例。原位杂交检测到IgM阳性细胞首先在孵化后18天出现在中华鳖脾脏中,随后在28天和64天的肾脏和肠中检测到,而IgD和IgY阳性细胞出现时间要晚于IgM。
Chinese soft-shelled turtle, Pelodiscus sinensis is one of important reptile species in aquaculture industry of China. The understanding of its immune system is not only important and necessary for developmental and comparative immunity in vertebrates, but also for aquacultural purpose. The present study aims to clone the cDNAs encoding the heavy chains and joining chain of IgM, IgD and IgY in Chinese soft-shelled turtle, and to analyze their expression at mRNA and protein levels. Furthermore, the ontogeny of IgM-, IgD- and IgY- producing cells in lymphoid tissues of the turtle was also analyzed. The cDNA sequence of J chain has been cloned from the Chinese soft-shelled turtle by reverse transcription (RT)-PCR and rapid amplification of cDNA ends (RACE). The cDNA sequence is 2,347 nt in length and contains an open reading frame of 480 nt encoding 160 aa including the signal sequence. The deduced amino acid sequence has a high degree of homology with that of an already reported turtle J chain (80.7%), and of chicken (71.3%), and a low degree of homology with those of amphibians (49.4% and 57.6%); but it is relatively low when compared with the sequence reported from skate (35.8%). It is thus implied that the J chain of P. sinensis is much more similar to avian, as also revealed in the phylogenetic tree. The expression of J chain was detected at 5 days past fertilization (dpf) by RT-PCR, and its expression increased following further developmental stages. The expression of J chain was detected in almost all organs examined at protein levels by Western blotting, except in fat tissue. The J chain-producing cells were detected in lymphoid-related tissues such as in spleen, kidney and intestine by using immunohistochemistry. By using real-time quantitative RT-PCR analysis, a significant up-regulation of J chain transcripts was observed in spleen, kidney and blood of the turtle injected with in-activated Aeromonas hydrophila T4, indicating the important immune role of J chain at early development and in response to bacterial infection.
Only three Ig isotypes, IgM, IgD, and IgA, were previously known in reptiles. Here, we describe IgM, IgD and a novel immunoglobulin heavy-chain isotypeυ(IgY) in Chinese soft-shelled turtle. The IgM and IgY genes of the turtle are characteristically similar to those described in other species, whereas IgD gene differs from IgD reported in other vertebrates. The turtle IgD contains 6 immunoglobulin heavy chain domains without evidence of intragenic duplications of exons as described in teleost IgD genes. The expression of IgM and IgD was detected early during embryonic development after fertilization, and their expression increased following further developmental stages. However, immunoglobulin Y could not be detected in spleen and kidney until 90 days after hatching out. Western blotting analysis demonstrated that the protein of IgM, IgD and IgY were expressed in almost all organs examined in adult turtles. The IgM-, IgD- and IgY- producing cells in lymphoid-related tissues such as in spleen, kidney and intestine, were investigated by using histochemistry. The transcriptional kinetics of the three Ig molecules in spleen, kidney and blood and antibody response kinetics in serum after bacterial pathogen stimulation were also analyzed by using real-time PCR and by using indirected enzyme-linked immunosorbent assay (ELISA), respectively. A significant up-regulation of J chain was observed at both mRNA and protein levels. Using flow cytometry, a higher proportion of IgM-pruducing cells was observed in spleen, peripheral blood and kidney when compared to proportions of IgD-and IgY- positive cells. The IgM-producing cells was firstly observed in spleen of P. sinensis at 18 dph, and then in kidney and intestine at 28 and 64 dph, respectively. However, IgD and IgY positive cells were detected much later.
引文
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