67kD层粘连蛋白受体及其调控通路在文蛤（Meretrix meretrix）细胞生长和幼虫发育中的功能解析
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摘要
层粘连蛋白受体（laminin receptor, LR）是存在于细胞表面的跨膜糖蛋白，是介导细胞与细胞间、细胞与细胞外基质间相互作用的一种非常重要的蛋白。LR通过与其配体层粘连蛋白（laminin, LN）的结合，构架了细胞同基膜间的信号传递通路，对胚胎早期发育、重要组织器官的形成以及维持生物体正常新陈代谢等生理活动中具有不可忽视的重要作用。目前，对层粘连蛋白受体以及其下游信号通路的研究多集中于模式动物，在贝类等低等物种中研究的很少。
文蛤（Meretrix meretrix）是我国重要的养殖贝类，也是海洋双壳贝类的重要代表物种之一。本论文以文蛤为实验对象，通过全长克隆获得了67kD LR基因（MmeLR）的全长cDNA序列，初步分析该基因的表达谱表明，其参与了文蛤的生长发育调控。在此基础上，进一步分别在细胞水平、幼虫阶段和成体阶段研究了MmeLR及其调控通路对文蛤生长发育的调控机制，主要研究结果详述如下：
通过RACE从文蛤中获得了MmeLR的全长cDNA序列。将MmeLR基因进行原核重组表达，得到带有GST标签的重组蛋白rMmeLR，制备了兔抗多克隆抗体。通过realtime PCR和western blot对MmeLR的组织分布特征进行了研究。利用far-western技术，验证了MmeLR及其配体层粘连蛋白（LN）间的相互作用，结果表明MmeLR和LN之间存在特异性相互作用。
通过优化培养条件，可以稳定培养文蛤血细胞、外套膜和肝胰脏等组织的原代细胞达2-3周，并使之维持较高活力，为在细胞水平上进行基因功能验证等打下了基础。在文蛤原代培养细胞中，进一步研究了MmeLR的功能。以matrigel（主要成分为LN）包被培养皿培养的文蛤外套膜细胞，在12h和24h时MmeLR的mRNA和蛋白表达量明显低于对照组，提示了LR在调控通路中的重要作用；同时，凋亡指示基因bcl-2和P53的mRNA表达量在试验组和对照组也有显著差异，进一步验证了MmeLR在细胞凋亡中的作用。
制备了MmeLR的双链RNA（dsRNA）和siRNA（small interference RNA）并在文蛤原代细胞中进行了干扰方法研究，沉默效率可以达到60%；制备了LR-pEGFP-N1和LR-pEGFP-ie1两种质粒，在人HeLa细胞系和昆虫SF9细胞系中进行了转染实验，结果显示可以稳定检测到EGFP阳性信号。在文蛤原代细胞中，对几种质粒进行了转移条件的探索，包括电穿孔法、脂质体转染法等。
在文蛤幼虫发育过程中，通过realtime PCR和western blot对MmeLR在文蛤幼虫不同发育阶段的表达谱进行了研究，使用免疫荧光技术分析了MmeLR在文蛤幼虫的时空分布，表明其参与了幼虫的发育。使用MmeLR的siRNA对文蛤幼虫进行了干扰实验，并对其下游调控通路进行了初步分析。对MmeLR下游分子钙调蛋白（MmeCAM）、钙调蛋白激酶IV（MmeCAMKIV）进行了全长克隆和表达谱研究，并使用CAM抑制剂三氟拉嗪（TFP）对MmeCaM在文蛤幼虫发育过程中的作用进行了分析。
以上研究部分阐明了LR基因及其调控通路在贝类细胞生长和幼虫发育中的作用，结合建立的贝类细胞原代培养技术、RNAi和基因过表达技术，将为海洋无脊椎动物功能基因的分析验证和功能解析提供新的研究平台。
Laminin receptor (LR) is a transmembrane glycoprotein expressed on the surfaceof the cell, which mediated interactions between cell and cell, cell and extracellularmatrix. The combination of LR and its ligand laminin (LN) conduct the signalingtransduction pathways between cell and extracellular matrix. It plays great roles inearly embryonic development, organogenesis and maintaining normal physiologicalactivities of organisms. At present, studies on LR and the signaling pathways itregulates mainly focused on the animal model, few studies on marine invertebrateswas reported.
The clam Meretrix meretrix is a representative species of marine bivalve, whichis an important commercial bivalve in China. In the present study, we cloned andcharacterized a67kD LR gene from M. meretrix and named it MmeLR. On this basis,further studies at the cellular level, larval and adult stages of the regulatorymechanisms of MmeLR and the pathways it regulates on the growth and developmentof M. meretrix were carried out.
A recombinant MmeLR (rMmeLR) protein was obtained by prokaryoticexpression. Its tissue expression characteristics were examined on both thetranscriptional and translational levels. The MmeLR mRNA and protein detected byrealtime PCR and western blots were primarily distributed in muscle tissues.Far-western analysis showed a specific interaction between rMmeLR and LN.
We successfully cultured primary cells from the different tissues of M. meretrixfor at least2weeks by a modified culture condition. The results of the binding assaysuggested a role of LR in cell adhesion and apoptosis in cultured primary cells ofmantle tissues from M. meretrix. The bcl-2mRNA expression levels in primary cells cultured in matrigel (mainly laminin) coated plates was significantly higher than incells cultured in non-coated plates at48h of culture, while the p53mRNA expressionpattern was inversely related to that of bcl-2, suggesting that MmeLR is involved inp53-dependent apoptosis, and the binding between MmeLR and laminin inhibitsapoptosis during primary cell culture.
Double strand RNA (dsRNA) and siRNA (small interference RNA) of MmeLRwere prepared to develop the method of RNAi in the primary cells of M. meretrix. Thesilencing efficiency was about60%. LR-pEGFP-N1and LR-pEGFP-ie1plasmidswere construted. Transfection experiments in HeLa and SF9cell lines were performedand EGFP positive signals can be stably detected. In M. meretrix primary cells, thetransfection conditions were explored, including electroporation, liposometransfection methods.
Realtime PCR, Western blot and immunofluorescence were used to analyze theexpression profile of MmeLR in different larval stages of M. meretrix. RNAi wasproformed on M. meretrix larvae using siRNA, and the downstream pathways wereanalyzed. The MmeLR downstream protein calmodulin (MmeCAM), calmodulinkinase IV (MmeCAMKIV) were cloned and analyzed. The CAM inhibitortrifluoperazine (TFP) was used to discover roles of MmeCaM in larval developmentof M. meretrix.
This study may increase the understanding of the role of LR in cell adhesion andapoptosis and help to improve the culture of primary cells of marine invertebrates.
文蛤（Meretrix meretrix）是我国重要的养殖贝类，也是海洋双壳贝类的重要代表物种之一。本论文以文蛤为实验对象，通过全长克隆获得了67kD LR基因（MmeLR）的全长cDNA序列，初步分析该基因的表达谱表明，其参与了文蛤的生长发育调控。在此基础上，进一步分别在细胞水平、幼虫阶段和成体阶段研究了MmeLR及其调控通路对文蛤生长发育的调控机制，主要研究结果详述如下：
通过RACE从文蛤中获得了MmeLR的全长cDNA序列。将MmeLR基因进行原核重组表达，得到带有GST标签的重组蛋白rMmeLR，制备了兔抗多克隆抗体。通过realtime PCR和western blot对MmeLR的组织分布特征进行了研究。利用far-western技术，验证了MmeLR及其配体层粘连蛋白（LN）间的相互作用，结果表明MmeLR和LN之间存在特异性相互作用。
通过优化培养条件，可以稳定培养文蛤血细胞、外套膜和肝胰脏等组织的原代细胞达2-3周，并使之维持较高活力，为在细胞水平上进行基因功能验证等打下了基础。在文蛤原代培养细胞中，进一步研究了MmeLR的功能。以matrigel（主要成分为LN）包被培养皿培养的文蛤外套膜细胞，在12h和24h时MmeLR的mRNA和蛋白表达量明显低于对照组，提示了LR在调控通路中的重要作用；同时，凋亡指示基因bcl-2和P53的mRNA表达量在试验组和对照组也有显著差异，进一步验证了MmeLR在细胞凋亡中的作用。
制备了MmeLR的双链RNA（dsRNA）和siRNA（small interference RNA）并在文蛤原代细胞中进行了干扰方法研究，沉默效率可以达到60%；制备了LR-pEGFP-N1和LR-pEGFP-ie1两种质粒，在人HeLa细胞系和昆虫SF9细胞系中进行了转染实验，结果显示可以稳定检测到EGFP阳性信号。在文蛤原代细胞中，对几种质粒进行了转移条件的探索，包括电穿孔法、脂质体转染法等。
在文蛤幼虫发育过程中，通过realtime PCR和western blot对MmeLR在文蛤幼虫不同发育阶段的表达谱进行了研究，使用免疫荧光技术分析了MmeLR在文蛤幼虫的时空分布，表明其参与了幼虫的发育。使用MmeLR的siRNA对文蛤幼虫进行了干扰实验，并对其下游调控通路进行了初步分析。对MmeLR下游分子钙调蛋白（MmeCAM）、钙调蛋白激酶IV（MmeCAMKIV）进行了全长克隆和表达谱研究，并使用CAM抑制剂三氟拉嗪（TFP）对MmeCaM在文蛤幼虫发育过程中的作用进行了分析。
以上研究部分阐明了LR基因及其调控通路在贝类细胞生长和幼虫发育中的作用，结合建立的贝类细胞原代培养技术、RNAi和基因过表达技术，将为海洋无脊椎动物功能基因的分析验证和功能解析提供新的研究平台。
Laminin receptor (LR) is a transmembrane glycoprotein expressed on the surfaceof the cell, which mediated interactions between cell and cell, cell and extracellularmatrix. The combination of LR and its ligand laminin (LN) conduct the signalingtransduction pathways between cell and extracellular matrix. It plays great roles inearly embryonic development, organogenesis and maintaining normal physiologicalactivities of organisms. At present, studies on LR and the signaling pathways itregulates mainly focused on the animal model, few studies on marine invertebrateswas reported.
The clam Meretrix meretrix is a representative species of marine bivalve, whichis an important commercial bivalve in China. In the present study, we cloned andcharacterized a67kD LR gene from M. meretrix and named it MmeLR. On this basis,further studies at the cellular level, larval and adult stages of the regulatorymechanisms of MmeLR and the pathways it regulates on the growth and developmentof M. meretrix were carried out.
A recombinant MmeLR (rMmeLR) protein was obtained by prokaryoticexpression. Its tissue expression characteristics were examined on both thetranscriptional and translational levels. The MmeLR mRNA and protein detected byrealtime PCR and western blots were primarily distributed in muscle tissues.Far-western analysis showed a specific interaction between rMmeLR and LN.
We successfully cultured primary cells from the different tissues of M. meretrixfor at least2weeks by a modified culture condition. The results of the binding assaysuggested a role of LR in cell adhesion and apoptosis in cultured primary cells ofmantle tissues from M. meretrix. The bcl-2mRNA expression levels in primary cells cultured in matrigel (mainly laminin) coated plates was significantly higher than incells cultured in non-coated plates at48h of culture, while the p53mRNA expressionpattern was inversely related to that of bcl-2, suggesting that MmeLR is involved inp53-dependent apoptosis, and the binding between MmeLR and laminin inhibitsapoptosis during primary cell culture.
Double strand RNA (dsRNA) and siRNA (small interference RNA) of MmeLRwere prepared to develop the method of RNAi in the primary cells of M. meretrix. Thesilencing efficiency was about60%. LR-pEGFP-N1and LR-pEGFP-ie1plasmidswere construted. Transfection experiments in HeLa and SF9cell lines were performedand EGFP positive signals can be stably detected. In M. meretrix primary cells, thetransfection conditions were explored, including electroporation, liposometransfection methods.
Realtime PCR, Western blot and immunofluorescence were used to analyze theexpression profile of MmeLR in different larval stages of M. meretrix. RNAi wasproformed on M. meretrix larvae using siRNA, and the downstream pathways wereanalyzed. The MmeLR downstream protein calmodulin (MmeCAM), calmodulinkinase IV (MmeCAMKIV) were cloned and analyzed. The CAM inhibitortrifluoperazine (TFP) was used to discover roles of MmeCaM in larval developmentof M. meretrix.
This study may increase the understanding of the role of LR in cell adhesion andapoptosis and help to improve the culture of primary cells of marine invertebrates.
引文
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