黄颡鱼IgM分子及其表达研究
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摘要
黄颡鱼(Pelteobagrus fulvidraco Richardson)是我国重要的淡水养殖鱼类,对其免疫系统的研究不仅在黄颡鱼及其它鱼类病害防治方面具有重要价值,而且在脊椎动物比较免疫及免疫发生方面也具有重要意义。本论文对编码黄颡鱼免疫球蛋白(Ig)M重链(H)和轻链(L)基因cDNA进行了克隆,并研究了IgM的表达规律及其传递和个体发生
运用RT和RACE PCR方法扩增出黄颡鱼IgM重链(ycIgM)和免疫球蛋白轻链(ycIgL) L1、L2和L3全长cDNA。 yclgM cDNA全长为1993bp,包含5'非编码区42bp,3'非编码区229bp,开放阅读框1722bp,编码573个氨基酸。ycIgL1cDNA全长为1021bp,包含5’非编码区39bp,3'非编码区259bp,开放阅读框723bp,编码240个氨基酸。ycIg L2cDNA全长为953bp,包含5'非编码区57bp,3'非编码区164bp,开放阅读框732bp,编码243个氨基酸。ycIg L3cDNA全长为965bp,包含5'非编码区58bp,3'非编码区184bp,开放阅读框723bp,编码240个氨基酸。序列比对分析表明ycIgM、ycIgL1ycIgL2和ycIgL3基因与斑点叉尾鯝的相似性最高为52.3%、90.%、80.2%和68.9%,并在进化树中分别与斑点叉尾鯝的IgM、IgL-G、IgL-2和IgL-F聚为一支。
RT-PCR表明yclgM基因主要在头肾、中肾、脾脏和血液中表达,而在心脏、肌肉和脑中表达量很少。ycIg L1基因主要在头肾、脾脏、鳃、肠和血液表达,在肌肉、皮肤和脑中表达量很少；L2基因主要在头肾、脾脏、鳃、肠和血液表达,在心脏和脑中表达量很少；L3基因主要在脾脏、鳃、肠、肝和血液中表达,而在心脏和脑中表达量很少。
Real-time PCR分析表明,注射嗜水气单胞菌后,ycIgM基因在黄颡鱼头肾、脾脏、肝和血液表达量在15d有显著上升,而在肠组织中5d内显著增加,15d恢复到刺激前水平。原位杂交分析表明,ycIgM基因阳性细胞出现在头肾、脾脏、鳃和肠组织中,注射嗜水气单胞菌15d后,头肾和脾脏组织中阳性细胞明显增加。而此时肝脏组织中也开始检测到了ycIgM基因阳性细胞。
通过原核表达载体pQE30,利用IPTG诱导,在大肠杆菌表达菌株M15获得大量可溶ycIgM蛋白。以纯化后的重组ycIgM蛋白为抗原,免疫新西兰兔制备了抗ycIgM蛋白的多克隆抗血清,ELISA测定抗血清效价约为1:512000; Western blot分析显示IgM在健康黄颡鱼头肾、脾脏、血清、皮肤、鳃、肠、肝和中肾中表达；免疫组化分析表明,ycIgM蛋白阳性信号出现在头肾、脾脏、鳃、肠和肝脏组织中,注射嗜水气单胞菌15d后,在这些组织中阳性信号有不同程度的变化。免疫荧光分析表明丰富的ycIgM蛋白荧光信号出现在头肾、脾脏、鳃、肠和肝脏组织中。
利用RT-PCR和Real-time PCR技术,在黄颡鱼卵和胚胎中检测到ycIgM mRNA。胚胎孵化后14d时仔鱼匀浆中检测到ycIgM mRNA,而在3-7d内没有检测到。利用Elisa技术,在黄颡鱼卵、胚胎、仔鱼各时间点都检测到yclgM蛋白,而且逐渐下降,在孵化9d蛋白表达量最低,到14d蛋白表达开始上升。这些研究表明,ycIgM mRNA和蛋白在仔鱼早期没有合成,存在于早期仔鱼的ycIgM蛋白来自于亲体。
Yellow catfish Pelteobagrus fulvidraco, as one of important commercial fishes in aquaculture industry of China, was threatened seriously by many kinds of diseases in recent years. It is thus important and necessary to understand developmental and comparative immunity in vertebrates and immune system in Yellow catfish for disease controlling. In this paper, the cDNAs encoding IgM heavy chain and light chains were cloned and analyzed, and expression pattern, maternal transfer and ontogeny of IgM were studied in yellow catfish.
cDNAs encoding IgM heavy chain (ycIgM) and light chains (ycIgL) were cloned using RACE-PCR and RT-PCR methods in P. fulvidraco. The full-length ycIgM gene cDNA contains1993nucleotides, including42nucleotides of5'-untranslated region (UTR),229nucleotides of3'-UTR, and1722nucleotides of the open reading frame encoding a putative protein of573amino acids; ycIgL1gene cDNA contains1021nucleotides, including39nucleotides of5'-UTR,259nucleotides of3'-UTR, and723nucleotides of the open reading frame encoding a putative protein of240amino acids; ycIgL2gene cDNA contains953nucleotides, including57nucleotides of5'-UTR,164nucleotides of3'-UTR, and732nucleotides of the open reading frame encoding a putative protein of243amino acids; ycIgL3gene cDNA contains965nucleotides, including58nucleotides of5'-UTR,184nucleotides of3'-UTR, and723nucleotides of the open reading frame encoding a putative protein of240amino acids. Compared with those of other teleost species ycIgM, L1, L2and L3shared the highest identity with that in channel catfish (52.3%,90.%,80.2%, and68.9%), respectively. Phylogenetic tree based on some teleost IgM heavy chain amino acids suggested IgM in P. fulvidraco was clustered closely with that of I. punctatus.
The abundance of ycIgM transcripts, was higher in the head kidney, kidney, blood cells and spleen than that in heart, muscle, and brain. Higher abundance of ycIgL1transcripts was found in the head kidney, spleen, gill, intestine and blood cells, whereas very little expression in the muscle, skin, and brain; Higher abundance of ycIgL2transcripts was found in the head kidney, spleen, gill, intestine and blood cells, whereas very little expression in the heart and brain; Higher abundance of ycIgL3transcripts was found in the spleen, gill, liver, intestine, and blood cells, whereas very little expression in the heart and brain.
The result of the real-time PCR also revealed that IgM mRNA expression increased significantly in blood cells, spleen, head kidney, liver, and intestine after injection of Aeromonas hydrophila. In situ hybridization results showed that ycIgM-producing cells were located in all the tissues investigated, such as head kidney, spleen and intestine, and increased in density in them post injection of A. hydrophila. This was the reason why the IgM mRNA expression increased significantly in the organs mentioned above after stimulation of the bacterial pathogen.
Using prokaryotic expression vector pQE30and induction of IPTG, the fusion protein of ycIgM heavy chain was produced in E. coli M15. Using the recombinant ycIgM protein as antigen, The anti-ycIgM polyclonal antibody was gotten by means of immuning a New Zealand white rabbit. The result of the Enzyme-linked Immunosorbent Assay (ELISA) showed that the highest titer of anti-serum was1:512000. Western blotting analysis showed IgM protein, about72kDa was detected in head kidney, spleen, serum, skin, intestine, gill, liver and kidney. Immuno-histochemical analysis showed that ycIgM protein positive signal appeared in head kidney, spleen, gill, intestine and liver tissue, and increased significantly at15d after infection of A. hydrophila.
Immunohistochemical and immuno-fluorescence analysis showed that the rich positive protein signals of ycIgM appear in head kidney, spleen, intestine, gill and liver tissue, and changeded significantly in the signals at15d after infection of A. hydrophila.
Using RT-PCR and Real-time PCR technology, ycIgM mRNA was detected in eggs, embryos and larva at14d after hatching, and wasn't detected in larva at3-7d after hatching. ELISA analysis showed that ycIgM protein was detected in eggs, embryos and larva in all phases. ycIgM protein decreased gradually in egg, embryos and early larva, and increased at14d after hatching. These studies showed synthesis of ycIgM mRNA and protein did not appear in embryos and early larva, and suggested that ycIgM proteins in them came from maternal yellow catfish.
运用RT和RACE PCR方法扩增出黄颡鱼IgM重链(ycIgM)和免疫球蛋白轻链(ycIgL) L1、L2和L3全长cDNA。 yclgM cDNA全长为1993bp,包含5'非编码区42bp,3'非编码区229bp,开放阅读框1722bp,编码573个氨基酸。ycIgL1cDNA全长为1021bp,包含5’非编码区39bp,3'非编码区259bp,开放阅读框723bp,编码240个氨基酸。ycIg L2cDNA全长为953bp,包含5'非编码区57bp,3'非编码区164bp,开放阅读框732bp,编码243个氨基酸。ycIg L3cDNA全长为965bp,包含5'非编码区58bp,3'非编码区184bp,开放阅读框723bp,编码240个氨基酸。序列比对分析表明ycIgM、ycIgL1ycIgL2和ycIgL3基因与斑点叉尾鯝的相似性最高为52.3%、90.%、80.2%和68.9%,并在进化树中分别与斑点叉尾鯝的IgM、IgL-G、IgL-2和IgL-F聚为一支。
RT-PCR表明yclgM基因主要在头肾、中肾、脾脏和血液中表达,而在心脏、肌肉和脑中表达量很少。ycIg L1基因主要在头肾、脾脏、鳃、肠和血液表达,在肌肉、皮肤和脑中表达量很少；L2基因主要在头肾、脾脏、鳃、肠和血液表达,在心脏和脑中表达量很少；L3基因主要在脾脏、鳃、肠、肝和血液中表达,而在心脏和脑中表达量很少。
Real-time PCR分析表明,注射嗜水气单胞菌后,ycIgM基因在黄颡鱼头肾、脾脏、肝和血液表达量在15d有显著上升,而在肠组织中5d内显著增加,15d恢复到刺激前水平。原位杂交分析表明,ycIgM基因阳性细胞出现在头肾、脾脏、鳃和肠组织中,注射嗜水气单胞菌15d后,头肾和脾脏组织中阳性细胞明显增加。而此时肝脏组织中也开始检测到了ycIgM基因阳性细胞。
通过原核表达载体pQE30,利用IPTG诱导,在大肠杆菌表达菌株M15获得大量可溶ycIgM蛋白。以纯化后的重组ycIgM蛋白为抗原,免疫新西兰兔制备了抗ycIgM蛋白的多克隆抗血清,ELISA测定抗血清效价约为1:512000; Western blot分析显示IgM在健康黄颡鱼头肾、脾脏、血清、皮肤、鳃、肠、肝和中肾中表达；免疫组化分析表明,ycIgM蛋白阳性信号出现在头肾、脾脏、鳃、肠和肝脏组织中,注射嗜水气单胞菌15d后,在这些组织中阳性信号有不同程度的变化。免疫荧光分析表明丰富的ycIgM蛋白荧光信号出现在头肾、脾脏、鳃、肠和肝脏组织中。
利用RT-PCR和Real-time PCR技术,在黄颡鱼卵和胚胎中检测到ycIgM mRNA。胚胎孵化后14d时仔鱼匀浆中检测到ycIgM mRNA,而在3-7d内没有检测到。利用Elisa技术,在黄颡鱼卵、胚胎、仔鱼各时间点都检测到yclgM蛋白,而且逐渐下降,在孵化9d蛋白表达量最低,到14d蛋白表达开始上升。这些研究表明,ycIgM mRNA和蛋白在仔鱼早期没有合成,存在于早期仔鱼的ycIgM蛋白来自于亲体。
Yellow catfish Pelteobagrus fulvidraco, as one of important commercial fishes in aquaculture industry of China, was threatened seriously by many kinds of diseases in recent years. It is thus important and necessary to understand developmental and comparative immunity in vertebrates and immune system in Yellow catfish for disease controlling. In this paper, the cDNAs encoding IgM heavy chain and light chains were cloned and analyzed, and expression pattern, maternal transfer and ontogeny of IgM were studied in yellow catfish.
cDNAs encoding IgM heavy chain (ycIgM) and light chains (ycIgL) were cloned using RACE-PCR and RT-PCR methods in P. fulvidraco. The full-length ycIgM gene cDNA contains1993nucleotides, including42nucleotides of5'-untranslated region (UTR),229nucleotides of3'-UTR, and1722nucleotides of the open reading frame encoding a putative protein of573amino acids; ycIgL1gene cDNA contains1021nucleotides, including39nucleotides of5'-UTR,259nucleotides of3'-UTR, and723nucleotides of the open reading frame encoding a putative protein of240amino acids; ycIgL2gene cDNA contains953nucleotides, including57nucleotides of5'-UTR,164nucleotides of3'-UTR, and732nucleotides of the open reading frame encoding a putative protein of243amino acids; ycIgL3gene cDNA contains965nucleotides, including58nucleotides of5'-UTR,184nucleotides of3'-UTR, and723nucleotides of the open reading frame encoding a putative protein of240amino acids. Compared with those of other teleost species ycIgM, L1, L2and L3shared the highest identity with that in channel catfish (52.3%,90.%,80.2%, and68.9%), respectively. Phylogenetic tree based on some teleost IgM heavy chain amino acids suggested IgM in P. fulvidraco was clustered closely with that of I. punctatus.
The abundance of ycIgM transcripts, was higher in the head kidney, kidney, blood cells and spleen than that in heart, muscle, and brain. Higher abundance of ycIgL1transcripts was found in the head kidney, spleen, gill, intestine and blood cells, whereas very little expression in the muscle, skin, and brain; Higher abundance of ycIgL2transcripts was found in the head kidney, spleen, gill, intestine and blood cells, whereas very little expression in the heart and brain; Higher abundance of ycIgL3transcripts was found in the spleen, gill, liver, intestine, and blood cells, whereas very little expression in the heart and brain.
The result of the real-time PCR also revealed that IgM mRNA expression increased significantly in blood cells, spleen, head kidney, liver, and intestine after injection of Aeromonas hydrophila. In situ hybridization results showed that ycIgM-producing cells were located in all the tissues investigated, such as head kidney, spleen and intestine, and increased in density in them post injection of A. hydrophila. This was the reason why the IgM mRNA expression increased significantly in the organs mentioned above after stimulation of the bacterial pathogen.
Using prokaryotic expression vector pQE30and induction of IPTG, the fusion protein of ycIgM heavy chain was produced in E. coli M15. Using the recombinant ycIgM protein as antigen, The anti-ycIgM polyclonal antibody was gotten by means of immuning a New Zealand white rabbit. The result of the Enzyme-linked Immunosorbent Assay (ELISA) showed that the highest titer of anti-serum was1:512000. Western blotting analysis showed IgM protein, about72kDa was detected in head kidney, spleen, serum, skin, intestine, gill, liver and kidney. Immuno-histochemical analysis showed that ycIgM protein positive signal appeared in head kidney, spleen, gill, intestine and liver tissue, and increased significantly at15d after infection of A. hydrophila.
Immunohistochemical and immuno-fluorescence analysis showed that the rich positive protein signals of ycIgM appear in head kidney, spleen, intestine, gill and liver tissue, and changeded significantly in the signals at15d after infection of A. hydrophila.
Using RT-PCR and Real-time PCR technology, ycIgM mRNA was detected in eggs, embryos and larva at14d after hatching, and wasn't detected in larva at3-7d after hatching. ELISA analysis showed that ycIgM protein was detected in eggs, embryos and larva in all phases. ycIgM protein decreased gradually in egg, embryos and early larva, and increased at14d after hatching. These studies showed synthesis of ycIgM mRNA and protein did not appear in embryos and early larva, and suggested that ycIgM proteins in them came from maternal yellow catfish.
引文
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