人E-钙粘素融合蛋白基质构建干细胞微环境的研究
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摘要
细胞与细胞、细胞与细胞外基质之间存在着复杂的联系,细胞表面的受体与细胞外微环境相互作用,决定了细胞的粘附、增殖、迁移、分化,以及特异性功能表达。近年来,组织工程和再生医学的研究取得了重要的进展,为各领域的研究拓宽了一条新的途径。利用生物材料、医用材料等仿生构建细胞外微环境的应用已经成为生物材料领域研究的重点,其中,以基因工程技术为基础的新型生物材料---融合蛋白的研制已成为设计和构建细胞外微环境生物材料研究的新方向。本论文通过生物合成人E-钙粘素融合蛋白,研究开发仿生细胞-细胞间相互作用的生物材料,并仿生构建细胞外微环境调控细胞行为,其不仅能有效推动基础生物学的研究,同时在组织工程和再生医学等领域也有广泛的应用。
首先,我们重组了人细胞间粘附分子E-钙粘蛋白(hE-cadherin)的胞外域与免疫球蛋白G(IgG)的Fc域,生物合成人E-钙粘素融合蛋白(hE-cadherin-Fc)。基因工程技术构建了融合蛋白表达载体,利用293F细胞表达体系进行了融合蛋白的表达并纯化,Elisa和western blotting检测表明我们成功制备了高纯度的hE-cadherin-Fc。进一步将融合蛋白应用于疏水材料的表面修饰,仿生构建了细胞-细胞间相互作用的生物材料细胞外微环境并用于人骨髓间充质干细胞(hMSCs)的增殖及定向分化研究。接触角(WCA)及原子力显微镜(AFM)检测表明：hE-cadherin-Fc能够利用Fc结构域稳定的吸附在疏水材料表面,同时改善了材料表面的形态及亲水性,形成稳定的细胞外基质。与组织细胞培养板(TC-PS)和明胶(Gelatin)基质相比,hE-cadherin-Fc基质能显著增强hMSCs的粘附,并促进细胞增殖；同时未分化性标志物CD105的流式细胞术鉴定显示：hMSCs保持了良好的未分化性。hE-cadherin-Fc基质对hMSCs生物学行为的调控及其分子机制的进一步研究,发现融合蛋白基质显著提高了E钙粘素信号通路的相关分子表达水平,初步揭示了在细胞培养初期hE-cadherin-Fc基质与细胞间的相互作用代替了细胞-细胞间的相互作用。
其次,我们考察了hE-cadherin-Fc基质对hMSCs向肝细胞定向诱导分化的影响。在hE-cadherin-Fc基质表面hMSCs向肝细胞诱导分化培养4周后,细胞形态、肝特异性标志物基因和蛋白的表达、肝细胞分化功能表达的检测表明：不同基质条件下hMSCs均有部分细胞分化成为肝细胞样细胞,与TC-PS和Gelatin基质相比,hE-cadherin-Fc基质表面分化的肝细胞样细胞表达肝前体细胞标志物CD117的阳性细胞比例高于TC-PS和Gelatin基质表面,并且肝细胞特异性标志物基因白蛋白(ALB)、细胞角蛋白18(CK18)和肝细胞核因子4(HNF4)的分子水平表达量更高；ALB蛋白免疫荧光染色结果也证明了ALB蛋白表达与其分子表达水平相同的结果。同时,肝分化功能指标糖原储存、吲哚青绿(ICG)摄取、白蛋白和尿素分泌的检测结果表明,hE-cadherin-Fc基质显著提高了人骨髓间充质干细胞源肝样细胞分化功能的表达。本研究结果表明hE-cadherin-Fc基质与相关液性因子协同作用促进hMSCs向肝细胞定向跨胚层分化的潜能,其分子机制还有待进一步深入研究,揭示了生物材料设计对其生物学功能的影响及应用前景。
最后,我们进一步探索了hE-cadherin-Fc用于纳米纤维支架和多孔支架的生物功能改性的研究。扫描电子显微镜(SEM)的结果显示,融合蛋白的修饰没有改变支架的三维微纳米结构。X射线光电子能谱(XPS)和Elisa检测结果表明,hE-cadherin-Fc在三维支架表面有效形成了稳定的融合蛋白层。WCA结果显示hE-cadherin-Fc表面修饰显著改善了三维支架表面的亲水性；同时,hE-cadherin-Fc基质化提高了hMSCs在三维支架上的粘附与增殖。细胞骨架染色和细胞形态结果表明,在hE-cadherin-Fc表面改性支架内的细胞更接近成纤维状。此外,茜素红染色表明hE-cadherin-Fc基质表面扩增的干细胞保持了良好的分化性能。本研究表明hE-cadherin-Fc可用于三维支架表面改性,提高支架与细胞的亲和性,进一步赋予支架生物功能,显示了其在干细胞三维环境构建及生物医学领域的应用前景。
综上所述,hE-cadherin-Fc融合蛋白可用于疏水材料的表面改性,由钙粘素的同源结合介导细胞与材料粘附,通过生物材料模拟细胞间结合仿生构建人工细胞外微环境,显著改善干细胞粘附与增殖,并促进干细胞定向分化调控,为生物材料在再生医学领域的应用开辟了新的思路。
Extracellular matrix (ECM), neighboring cells and soluble factors are most important effectors of cell behavior and function. Among them, cell-matrix interaction plays a fundamental role in regulating cell adhesion, migration, proliferation, differentiation and the expression of specific functions. In recent years, there was some important progress in tissue engineering and regenerative medicine, which widened a new theory in various research fields. The recombinant fusion proteins, an engineered artificial ECM, which was based on gene engineering technology, has gained extensive attention because of its selectivity to specific cell lineages and tissue responses. In this study, human E-cadherin-Fc fusion protein was constructed and used as a cell-cell adhesion biomimicking matrix for human mesenchymal stem cells (hMSCs). It can not only be used in basic biological research, but also in tissue engineering and regenerative medicine.
Firstly, a fusion protein consisting of human E-cadherin extracellular domain and the immunoglobulin G Fc region (hE-cadherin-Fc) was biosynthesized by gene engineering technology. The hE-cadherin-Fc fusion protein expressed and purified by the Free-Style MAX293Expression System was analyzed by Elisa and western blotting. Water contact angles (WCA) and Atomic Force Microscope (AFM) showed that the hE-cadherin-Fc was stably immobilized onto a polystyrene plate due to the hydrophobicity of the Fc domain, enhancing the surface wettability and topography. The hE-cadherin-Fc matrix markedly promoted the cell adhesion and proliferation of hMSCs comparing with the tissue culture-treated plate (TC-PS) and the gelatin-coated plate. Furthermore, the expanded hMSCs on the hE-cadherin-Fc were positive for CD105, showing the hE-cadherin-Fc matrix could maintain the undifferentiation of hMSCs. Additionally, the expressions of E-cadherin and β-catenin in the hMSCs were improved on the hE-cadherin-Fc matrix, suggesting that the cell-cell adhesion junctions were substituted by the interactions between the hE-cadherin-Fc matrix and the hMSCs during the initial culture stage in the absence of cell-cell interactions.
Secondly, we focused on the effects of the hE-cadherin-Fc matrix on the hepatic differentiation from hMSCs. After4weeks of directly differentiation on modified surfaces, hepatocyte-like cells were identified through cell morphology, RT-PCR, flow cytometry, immunofluorescence, Periodic acid-Shiff staining, indocyanine green (ICG) uptake and hepatocellular synthesis and metabolism functions by albumin/urea assays. Compared with the TC-PS and gelatin-coated surface, the expression of hepatic lineage surface markers CD117was higher on hE-cadherin-Fc surface. The mRNA expressions of hepatocyte-specific markers, such as ALB, CK18and HNF4, were performed by RT-PCR. The expression levels of these markers were found to be significantly upregulated during the differentiation on hE-cadherin-Fc surface. Similarly, the expression of ALB protein which was detected by immunofluorescence staining appeared to be higher than TC-PS and gelatin-coated surface. To confirm whether differentiated hepatocyte-like cells derived from hMSC were functionally competent, we examined the glycogen storage, ICG uptake, albumin secretion and urea production. It demonstrated that the differentiated cells expressed higher function on hE-cadherin-Fc-coated surface. These results show that the hE-cadherin-Fc may be a promising artificial extracellular matrix (ECM) for the hMSCs differentiation via the homophilic interaction of hE-cadherin and the synergy between cell adhesion molecular and growth factors.
Lastly, we fabricated nanofibrous scaffolds and porous scaffolds, modified with hE-cadherin-Fc fusion protein and studied the synergistic effect of the scaffolds. The morphology of three-dimensional scaffolds which modified by hE-cadherin-Fc fusion protein was not changed by using scanning electron microscopy (SEM). Surface modification was investigated via monitoring the changes of the surface by X-ray photoelectron spectroscopy (XPS) and Elisa showed that the hE-cadherin-Fc was stably modified on three-dimensional scaffolds. WCA measurements confirmed that the hE-cadherin-Fc fusion protein could improve the surface properties of scaffolds. The hE-cadherin-Fc matrix markedly promoted the cell adhesion and proliferation of hMSCs on three-dimensional scaffolds. The cell morphology was evaluated by F-actin staining, and the cells had a more spreading morphology on hE-cadherin-Fc-coated scaffolds. The results demonstrated that the hE-cadherin-Fc improved the cell compatibility and biological functions of three-dimensional scaffolds.
In summary, hE-cadherin-Fc was successfully prepared as a cell-cell adhesion biomimicking matrix which can be used for the surface modification of hydrophobic materials effectively. The hE-cadherin-Fc fusion protein matrix could enhance the adhesion and proliferation of the hMSCs, and improve the differentiation regulation of stem cells. The hE-cadherin-Fc was shown to be a promising artificial ECM for hMSCs.
首先,我们重组了人细胞间粘附分子E-钙粘蛋白(hE-cadherin)的胞外域与免疫球蛋白G(IgG)的Fc域,生物合成人E-钙粘素融合蛋白(hE-cadherin-Fc)。基因工程技术构建了融合蛋白表达载体,利用293F细胞表达体系进行了融合蛋白的表达并纯化,Elisa和western blotting检测表明我们成功制备了高纯度的hE-cadherin-Fc。进一步将融合蛋白应用于疏水材料的表面修饰,仿生构建了细胞-细胞间相互作用的生物材料细胞外微环境并用于人骨髓间充质干细胞(hMSCs)的增殖及定向分化研究。接触角(WCA)及原子力显微镜(AFM)检测表明：hE-cadherin-Fc能够利用Fc结构域稳定的吸附在疏水材料表面,同时改善了材料表面的形态及亲水性,形成稳定的细胞外基质。与组织细胞培养板(TC-PS)和明胶(Gelatin)基质相比,hE-cadherin-Fc基质能显著增强hMSCs的粘附,并促进细胞增殖；同时未分化性标志物CD105的流式细胞术鉴定显示：hMSCs保持了良好的未分化性。hE-cadherin-Fc基质对hMSCs生物学行为的调控及其分子机制的进一步研究,发现融合蛋白基质显著提高了E钙粘素信号通路的相关分子表达水平,初步揭示了在细胞培养初期hE-cadherin-Fc基质与细胞间的相互作用代替了细胞-细胞间的相互作用。
其次,我们考察了hE-cadherin-Fc基质对hMSCs向肝细胞定向诱导分化的影响。在hE-cadherin-Fc基质表面hMSCs向肝细胞诱导分化培养4周后,细胞形态、肝特异性标志物基因和蛋白的表达、肝细胞分化功能表达的检测表明：不同基质条件下hMSCs均有部分细胞分化成为肝细胞样细胞,与TC-PS和Gelatin基质相比,hE-cadherin-Fc基质表面分化的肝细胞样细胞表达肝前体细胞标志物CD117的阳性细胞比例高于TC-PS和Gelatin基质表面,并且肝细胞特异性标志物基因白蛋白(ALB)、细胞角蛋白18(CK18)和肝细胞核因子4(HNF4)的分子水平表达量更高；ALB蛋白免疫荧光染色结果也证明了ALB蛋白表达与其分子表达水平相同的结果。同时,肝分化功能指标糖原储存、吲哚青绿(ICG)摄取、白蛋白和尿素分泌的检测结果表明,hE-cadherin-Fc基质显著提高了人骨髓间充质干细胞源肝样细胞分化功能的表达。本研究结果表明hE-cadherin-Fc基质与相关液性因子协同作用促进hMSCs向肝细胞定向跨胚层分化的潜能,其分子机制还有待进一步深入研究,揭示了生物材料设计对其生物学功能的影响及应用前景。
最后,我们进一步探索了hE-cadherin-Fc用于纳米纤维支架和多孔支架的生物功能改性的研究。扫描电子显微镜(SEM)的结果显示,融合蛋白的修饰没有改变支架的三维微纳米结构。X射线光电子能谱(XPS)和Elisa检测结果表明,hE-cadherin-Fc在三维支架表面有效形成了稳定的融合蛋白层。WCA结果显示hE-cadherin-Fc表面修饰显著改善了三维支架表面的亲水性；同时,hE-cadherin-Fc基质化提高了hMSCs在三维支架上的粘附与增殖。细胞骨架染色和细胞形态结果表明,在hE-cadherin-Fc表面改性支架内的细胞更接近成纤维状。此外,茜素红染色表明hE-cadherin-Fc基质表面扩增的干细胞保持了良好的分化性能。本研究表明hE-cadherin-Fc可用于三维支架表面改性,提高支架与细胞的亲和性,进一步赋予支架生物功能,显示了其在干细胞三维环境构建及生物医学领域的应用前景。
综上所述,hE-cadherin-Fc融合蛋白可用于疏水材料的表面改性,由钙粘素的同源结合介导细胞与材料粘附,通过生物材料模拟细胞间结合仿生构建人工细胞外微环境,显著改善干细胞粘附与增殖,并促进干细胞定向分化调控,为生物材料在再生医学领域的应用开辟了新的思路。
Extracellular matrix (ECM), neighboring cells and soluble factors are most important effectors of cell behavior and function. Among them, cell-matrix interaction plays a fundamental role in regulating cell adhesion, migration, proliferation, differentiation and the expression of specific functions. In recent years, there was some important progress in tissue engineering and regenerative medicine, which widened a new theory in various research fields. The recombinant fusion proteins, an engineered artificial ECM, which was based on gene engineering technology, has gained extensive attention because of its selectivity to specific cell lineages and tissue responses. In this study, human E-cadherin-Fc fusion protein was constructed and used as a cell-cell adhesion biomimicking matrix for human mesenchymal stem cells (hMSCs). It can not only be used in basic biological research, but also in tissue engineering and regenerative medicine.
Firstly, a fusion protein consisting of human E-cadherin extracellular domain and the immunoglobulin G Fc region (hE-cadherin-Fc) was biosynthesized by gene engineering technology. The hE-cadherin-Fc fusion protein expressed and purified by the Free-Style MAX293Expression System was analyzed by Elisa and western blotting. Water contact angles (WCA) and Atomic Force Microscope (AFM) showed that the hE-cadherin-Fc was stably immobilized onto a polystyrene plate due to the hydrophobicity of the Fc domain, enhancing the surface wettability and topography. The hE-cadherin-Fc matrix markedly promoted the cell adhesion and proliferation of hMSCs comparing with the tissue culture-treated plate (TC-PS) and the gelatin-coated plate. Furthermore, the expanded hMSCs on the hE-cadherin-Fc were positive for CD105, showing the hE-cadherin-Fc matrix could maintain the undifferentiation of hMSCs. Additionally, the expressions of E-cadherin and β-catenin in the hMSCs were improved on the hE-cadherin-Fc matrix, suggesting that the cell-cell adhesion junctions were substituted by the interactions between the hE-cadherin-Fc matrix and the hMSCs during the initial culture stage in the absence of cell-cell interactions.
Secondly, we focused on the effects of the hE-cadherin-Fc matrix on the hepatic differentiation from hMSCs. After4weeks of directly differentiation on modified surfaces, hepatocyte-like cells were identified through cell morphology, RT-PCR, flow cytometry, immunofluorescence, Periodic acid-Shiff staining, indocyanine green (ICG) uptake and hepatocellular synthesis and metabolism functions by albumin/urea assays. Compared with the TC-PS and gelatin-coated surface, the expression of hepatic lineage surface markers CD117was higher on hE-cadherin-Fc surface. The mRNA expressions of hepatocyte-specific markers, such as ALB, CK18and HNF4, were performed by RT-PCR. The expression levels of these markers were found to be significantly upregulated during the differentiation on hE-cadherin-Fc surface. Similarly, the expression of ALB protein which was detected by immunofluorescence staining appeared to be higher than TC-PS and gelatin-coated surface. To confirm whether differentiated hepatocyte-like cells derived from hMSC were functionally competent, we examined the glycogen storage, ICG uptake, albumin secretion and urea production. It demonstrated that the differentiated cells expressed higher function on hE-cadherin-Fc-coated surface. These results show that the hE-cadherin-Fc may be a promising artificial extracellular matrix (ECM) for the hMSCs differentiation via the homophilic interaction of hE-cadherin and the synergy between cell adhesion molecular and growth factors.
Lastly, we fabricated nanofibrous scaffolds and porous scaffolds, modified with hE-cadherin-Fc fusion protein and studied the synergistic effect of the scaffolds. The morphology of three-dimensional scaffolds which modified by hE-cadherin-Fc fusion protein was not changed by using scanning electron microscopy (SEM). Surface modification was investigated via monitoring the changes of the surface by X-ray photoelectron spectroscopy (XPS) and Elisa showed that the hE-cadherin-Fc was stably modified on three-dimensional scaffolds. WCA measurements confirmed that the hE-cadherin-Fc fusion protein could improve the surface properties of scaffolds. The hE-cadherin-Fc matrix markedly promoted the cell adhesion and proliferation of hMSCs on three-dimensional scaffolds. The cell morphology was evaluated by F-actin staining, and the cells had a more spreading morphology on hE-cadherin-Fc-coated scaffolds. The results demonstrated that the hE-cadherin-Fc improved the cell compatibility and biological functions of three-dimensional scaffolds.
In summary, hE-cadherin-Fc was successfully prepared as a cell-cell adhesion biomimicking matrix which can be used for the surface modification of hydrophobic materials effectively. The hE-cadherin-Fc fusion protein matrix could enhance the adhesion and proliferation of the hMSCs, and improve the differentiation regulation of stem cells. The hE-cadherin-Fc was shown to be a promising artificial ECM for hMSCs.
引文
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