海马放射状胶质细胞的体外诱导激活及向胆碱能神经元的分化
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摘要
第一部分：海马放射状胶质细胞的体外诱导激活及其神经干细胞样特性
目的探讨海马放射状胶质细胞在切割穹窿海马伞去神经支配海马提取液诱导下的形态学变化以及细胞增殖、胚胎源性和向神经元及胶质细胞分化的神经干细胞样特性。
方法取生后1 d的SD大鼠海马组织,应用差速贴壁及摇床振荡法分离纯化海马放射状胶质细胞。将纯化的海马放射状胶质细胞接种于24孔培养板中,分成切割组和正常组,切割组加入含5%(v/v)切割穹窿海马伞去神经支配海马提取液的DMEM/F12培养液,正常组加入含5%(v/v)正常侧海马提取液的DMEM/F12培养液,同时在两组中加入5-溴-2-脱氧尿嘧啶(BrdU)标记增殖的细胞,并分别于培养后1、3、7和14 d时行BLBP免疫荧光检测和Hoechst标记；同时用BLBP/BrdU、BLBP/nestin、BLBP/MAP-2、BLBP/GFAP、BLBP/CNP免疫荧光双标技术观察其增殖特性和胚胎源性以及向神经元、星型胶质细胞、少突胶质细胞分化的情况,检测BrdU阳性细胞占BLBP阳性细胞的百分比、BLBP阳性细胞的周长和面积,以及两组中BLBP/nestin、BLBP/MAP-2、BLBP/GFAP、BLBP/CNP双标细胞占BLBP阳性细胞的百分比。应用Stata10.0统计软件行组间比较。
结果BLBP免疫荧光检测结果显示,通过上述的纯化培养方法,我们获得了几近100%的海马放射状胶质细胞,BLBP在细胞浆、细胞核及突起中均有表达。切割组BrdU阳性细胞占BLBP阳性细胞的百分比明显高于正常组(切割组：56.86±8.52%；正常组：31.11±4.28%,P<0.01)。接种1 d后,两组细胞的胞体均较小,突起较短较细；3 d时,与正常组相比,切割组BLBP阳性细胞的胞体稍变大,突起稍变得粗而长；7 d时,与正常组相比,切割组BLBP阳性细胞的胞体明显变大,突起明显变粗、变长且交织成网状；14 d时两组细胞的胞体和突起均开始变细变短,正常组较为明显。两组各d BLBP阳性细胞的周长和面积的统计分析结果表明除1 d时两组BLBP阳性细胞的周长和面积无明显差异(P>0.05)外,其余各d切割组BLBP阳性细胞的周长和面积均高于正常组(P<0.01)。切割组nestin阳性细胞占BLBP阳性细胞的百分比也明显高于正常组(切割组：57.92±17.93%；正常组：23.26±9.85%,P<0.01)。并可见切割组中较多的BLBP阳性细胞向MAP-2阳性的神经元分化(切割组：46.13±14.92%；正常组：29.13±10.07%,P<0.05),虽然两组向GFAP阳性的星型胶质细胞和CNP阳性的少突胶质细胞分化在数量上无明显差异,但切割组中两种胶质细胞的胞体明显增大、突起明显丰富。
结论切割穹窿海马伞去神经支配海马提取液不仅可以明显诱导BLBP阳性放射状胶质细胞增殖、胞体明显增大、突起变粗变长呈“激活”状态,具胚胎源性,而且还可使其向神经元、星型胶质细胞和少突胶质细胞分化,表现为神经干细胞样特性。
第二部分：去神经支配海马提取液诱导海马放射状胶质细胞向胆碱能神经元的分化
目的在体外细胞培养中加入切割穹窿海马伞去神经支配海马提取液,模拟在体海马神经再生微环境,观察海马放射状胶质细胞向胆碱能神经元分化的情况。
方法将纯化的海马放射状胶质细胞分成切割组和正常组,切割组加入含5%(v/v)切割穹窿海马伞去神经支配海马提取液的DMEM/F12培养液,正常组加入含5%(v/v)正常侧海马提取液的DMEM/F12培养液,分别在培养7d后应用BLBP/ChAT免疫荧光双标、Real-time PCR及Western blot等技术观察两组放射状胶质细胞向胆碱能神经元的分化,以及表达ChAT mRNA和蛋白的情况。
结果ChAT免疫荧光结果显示,加入切割穹窿海马伞侧海马提取液后,与正常组相比,切割组中ChAT阳性的胆碱能神经元占BLBP阳性细胞的比例(41.62±9.97%)明显多于正常组(16.08±7.31%) (P<0.01),且切割组中ChAT阳性的胆碱能神经元的胞体较正常组明显增大,突起明显增粗增长。切割组ChAT mRNA水平明显高于正常组,约是正常组的5倍(P<0.01)。ChAT蛋白的表达量虽较低,但切割组(0.1141±0.0380)仍高于正常组(0.0423±0.0106)(P<0.05)。
结论切割穹窿海马伞侧海马提取液可明显诱导BLBP阳性放射状胶质细胞向胆碱能神经元分化。
Part I THE INDUCED ACTIVATION AND THE NEURAL STEM CELLS'CHARACTERISTICS OF HIPPOCAMPAL RADIAL GLIAL CELLS IN VITRO
Objective To explore the changes of hippocampal radial glial cells (RGCs) in the morphology, proliferation, embryogenicity and the neural stem cells (NSCs) identity towards to neurons and glial cells after the denervated fimbria-fornix transected hippocampal extracts induced.
Methods On the postnatal 1 day, the hippocampi of SD rats were acquired. RGCs were separated and purified by differential velocity adherent method and shakingbed oscillation technique. The purified RGCs were plated into 24-well plates and then divided into transection group and normal group. The DMEM/F12 medium containing 5% (vol/vol) fimbria-fornix transected hippocampal extracts was added into the transection group, while the normal group added the the DMEM/F12 medium containing 5% (vol/vol) normal hippocampal extracts. At the same time, BrdU were added into both groups to label the proliferated cells, the cells of both groups were detected by BLBP immuno-fluorescence and marked by Hoechst on the 1st,3rd,7th and 14th d respectively. Double-labeling immunofluorescence of BLBP/BrdU、BLBP/nestin、BLBP/MAP-2、BLBP/GFAP、BLBP/CNP were used to investigate the characteristics of proliferation, embryogenicity and the cell differentiation into neurons, astrocytes and oligodendrocytes. The percentage of BrdU positive cells over the BLBP positive cells, the perimeter and area of BLBP positive cells and the percentage of BLBP/nestin, BLBP/MAP-2, BLBP/GFAP, BLBP/CNP Double-labeling cells over the BLBP positive cells in two groups were deteced. Comparing analysis between both groups were analyzed by Statal0.0 statistical software.
Results Immunofluorescence assay of BLBP showed that we acquired nearly 100% RGCs by our purified method. BLBP expressed in the cytoplasm, nuclei and processes. The percentage of BrdU positive cells in the transection group was higher than the normal group (the transection group:56.86±8.52%; the normal group:31.11±4.28%; P< 0.01). On the 1 day after cell seeding, the cell bodies were small in both groups, the processes of both groups were also short and thin; On the 3rd day, compared to the normal group, the cell bodies of the transection group were slightly larger and the processes were also longer and thicker; On the 7th day, the cell bodies of RGCs in the transection group were significantly larger than the normal group, and the cell processes became much more longer and thicker, and interlaced into networks; on the 14th day, the cell bodies of both groups decreased slightly, and the processes also became shorter and thinner slightly. The statistical analysis of two groups'BLBP positive cells in the perimeter and area showed that the transection group is significant higher than the normal group (P<0.01) except the 1st day (P> 0.05). The percentage of nestin positive cells over the BLBP positive cells in the transection group was also higher than the normal group (the transection group:57.92±17.93%; the normal group: 23.26±9.85%; P< 0.01). And we observed more RGCs differentiated into MAP-2 positive neurons in the transection group (the transection group:46.13±14.92%; the normal group:29.13±10.07%; P< 0.01). Although there was no significant difference in the cell numbers between two groups when they differentiated into GFAP positive astrocytes and CNP positive oligodendrocyte, the cell bodies in the transection group were larger and the processes were also longer and thicker than the normal group.
Conclusions The results indicate that the denervated fimbria-fornix transected hippocampal extracts can significantly promote the proliferation of RGCs, enlarge the cell bodies, thicken and elongate the processes of BLBP positive RGCs which presenting the state of "activation" and containing the embryogenicity, and also can make RGCs present the identity of neural stem cells and differentiate into neurons, astrocytes and oligodendrocytes.
PartⅡTHE DIFFERENTION OF HIPPOCAMPAL RADIAL GLIAL CELLS INTO CHOLINERGIC NEURONS IN VITRO BY DENERVATED HIPPOCAMPAL EXTRACTS
Objective Fimbria-fornix transected hippocampal extracts were added into the cell culture medium to simulate the hippocampal internal environment in vitro, and to observe the situation of the RGCs differentiation into cholinergic neurons.
Methods Purified RGCs were divided into transection group and normal group. The DMEM/F12 medium supplemented with 5%(vol/vol) fimbria-fornix transected hippocampal extracts was added into the transection group, while the normal group the DMEM/F12 medium contains 5%(vol/vol) normal hippocampal extracts. BLBP/ChAT double-labeling immunofluorescence, Real-time PCR, and Western blot were used to observe the differentiation into cholinergic neurons, the levels of ChAT mRNA and protien in RGCs after 7 day inoculation.
Results Immunofluorescence assay of ChAT showed that the percentage of ChAT positive cholinergic neurons over the BLBP positive cells in the transection group(41.62±9.97%) was higher than the normal group(16.08±7.31%)(P< 0.01), and the cholinergic neuron bodies of transection group were larger and the processes were also longer and thicker than the normal group. The level of ChAT mRNA in the transection group was 5 times higher than the normal group (P< 0.01). The relative amount of ChAT protein in the transection group(0.1141±0.0380)was significant higher than that in the normal group(0.0423±0.0106)(P< 0.05), although both groups had low expression.
Conclusions The results indicate that the denervated hippocampal extracts after fimbria-fornix transection can significantly promote BLBP positive RGCs differentiate into cholinergic neurons.
目的探讨海马放射状胶质细胞在切割穹窿海马伞去神经支配海马提取液诱导下的形态学变化以及细胞增殖、胚胎源性和向神经元及胶质细胞分化的神经干细胞样特性。
方法取生后1 d的SD大鼠海马组织,应用差速贴壁及摇床振荡法分离纯化海马放射状胶质细胞。将纯化的海马放射状胶质细胞接种于24孔培养板中,分成切割组和正常组,切割组加入含5%(v/v)切割穹窿海马伞去神经支配海马提取液的DMEM/F12培养液,正常组加入含5%(v/v)正常侧海马提取液的DMEM/F12培养液,同时在两组中加入5-溴-2-脱氧尿嘧啶(BrdU)标记增殖的细胞,并分别于培养后1、3、7和14 d时行BLBP免疫荧光检测和Hoechst标记；同时用BLBP/BrdU、BLBP/nestin、BLBP/MAP-2、BLBP/GFAP、BLBP/CNP免疫荧光双标技术观察其增殖特性和胚胎源性以及向神经元、星型胶质细胞、少突胶质细胞分化的情况,检测BrdU阳性细胞占BLBP阳性细胞的百分比、BLBP阳性细胞的周长和面积,以及两组中BLBP/nestin、BLBP/MAP-2、BLBP/GFAP、BLBP/CNP双标细胞占BLBP阳性细胞的百分比。应用Stata10.0统计软件行组间比较。
结果BLBP免疫荧光检测结果显示,通过上述的纯化培养方法,我们获得了几近100%的海马放射状胶质细胞,BLBP在细胞浆、细胞核及突起中均有表达。切割组BrdU阳性细胞占BLBP阳性细胞的百分比明显高于正常组(切割组：56.86±8.52%；正常组：31.11±4.28%,P<0.01)。接种1 d后,两组细胞的胞体均较小,突起较短较细；3 d时,与正常组相比,切割组BLBP阳性细胞的胞体稍变大,突起稍变得粗而长；7 d时,与正常组相比,切割组BLBP阳性细胞的胞体明显变大,突起明显变粗、变长且交织成网状；14 d时两组细胞的胞体和突起均开始变细变短,正常组较为明显。两组各d BLBP阳性细胞的周长和面积的统计分析结果表明除1 d时两组BLBP阳性细胞的周长和面积无明显差异(P>0.05)外,其余各d切割组BLBP阳性细胞的周长和面积均高于正常组(P<0.01)。切割组nestin阳性细胞占BLBP阳性细胞的百分比也明显高于正常组(切割组：57.92±17.93%；正常组：23.26±9.85%,P<0.01)。并可见切割组中较多的BLBP阳性细胞向MAP-2阳性的神经元分化(切割组：46.13±14.92%；正常组：29.13±10.07%,P<0.05),虽然两组向GFAP阳性的星型胶质细胞和CNP阳性的少突胶质细胞分化在数量上无明显差异,但切割组中两种胶质细胞的胞体明显增大、突起明显丰富。
结论切割穹窿海马伞去神经支配海马提取液不仅可以明显诱导BLBP阳性放射状胶质细胞增殖、胞体明显增大、突起变粗变长呈“激活”状态,具胚胎源性,而且还可使其向神经元、星型胶质细胞和少突胶质细胞分化,表现为神经干细胞样特性。
第二部分：去神经支配海马提取液诱导海马放射状胶质细胞向胆碱能神经元的分化
目的在体外细胞培养中加入切割穹窿海马伞去神经支配海马提取液,模拟在体海马神经再生微环境,观察海马放射状胶质细胞向胆碱能神经元分化的情况。
方法将纯化的海马放射状胶质细胞分成切割组和正常组,切割组加入含5%(v/v)切割穹窿海马伞去神经支配海马提取液的DMEM/F12培养液,正常组加入含5%(v/v)正常侧海马提取液的DMEM/F12培养液,分别在培养7d后应用BLBP/ChAT免疫荧光双标、Real-time PCR及Western blot等技术观察两组放射状胶质细胞向胆碱能神经元的分化,以及表达ChAT mRNA和蛋白的情况。
结果ChAT免疫荧光结果显示,加入切割穹窿海马伞侧海马提取液后,与正常组相比,切割组中ChAT阳性的胆碱能神经元占BLBP阳性细胞的比例(41.62±9.97%)明显多于正常组(16.08±7.31%) (P<0.01),且切割组中ChAT阳性的胆碱能神经元的胞体较正常组明显增大,突起明显增粗增长。切割组ChAT mRNA水平明显高于正常组,约是正常组的5倍(P<0.01)。ChAT蛋白的表达量虽较低,但切割组(0.1141±0.0380)仍高于正常组(0.0423±0.0106)(P<0.05)。
结论切割穹窿海马伞侧海马提取液可明显诱导BLBP阳性放射状胶质细胞向胆碱能神经元分化。
Part I THE INDUCED ACTIVATION AND THE NEURAL STEM CELLS'CHARACTERISTICS OF HIPPOCAMPAL RADIAL GLIAL CELLS IN VITRO
Objective To explore the changes of hippocampal radial glial cells (RGCs) in the morphology, proliferation, embryogenicity and the neural stem cells (NSCs) identity towards to neurons and glial cells after the denervated fimbria-fornix transected hippocampal extracts induced.
Methods On the postnatal 1 day, the hippocampi of SD rats were acquired. RGCs were separated and purified by differential velocity adherent method and shakingbed oscillation technique. The purified RGCs were plated into 24-well plates and then divided into transection group and normal group. The DMEM/F12 medium containing 5% (vol/vol) fimbria-fornix transected hippocampal extracts was added into the transection group, while the normal group added the the DMEM/F12 medium containing 5% (vol/vol) normal hippocampal extracts. At the same time, BrdU were added into both groups to label the proliferated cells, the cells of both groups were detected by BLBP immuno-fluorescence and marked by Hoechst on the 1st,3rd,7th and 14th d respectively. Double-labeling immunofluorescence of BLBP/BrdU、BLBP/nestin、BLBP/MAP-2、BLBP/GFAP、BLBP/CNP were used to investigate the characteristics of proliferation, embryogenicity and the cell differentiation into neurons, astrocytes and oligodendrocytes. The percentage of BrdU positive cells over the BLBP positive cells, the perimeter and area of BLBP positive cells and the percentage of BLBP/nestin, BLBP/MAP-2, BLBP/GFAP, BLBP/CNP Double-labeling cells over the BLBP positive cells in two groups were deteced. Comparing analysis between both groups were analyzed by Statal0.0 statistical software.
Results Immunofluorescence assay of BLBP showed that we acquired nearly 100% RGCs by our purified method. BLBP expressed in the cytoplasm, nuclei and processes. The percentage of BrdU positive cells in the transection group was higher than the normal group (the transection group:56.86±8.52%; the normal group:31.11±4.28%; P< 0.01). On the 1 day after cell seeding, the cell bodies were small in both groups, the processes of both groups were also short and thin; On the 3rd day, compared to the normal group, the cell bodies of the transection group were slightly larger and the processes were also longer and thicker; On the 7th day, the cell bodies of RGCs in the transection group were significantly larger than the normal group, and the cell processes became much more longer and thicker, and interlaced into networks; on the 14th day, the cell bodies of both groups decreased slightly, and the processes also became shorter and thinner slightly. The statistical analysis of two groups'BLBP positive cells in the perimeter and area showed that the transection group is significant higher than the normal group (P<0.01) except the 1st day (P> 0.05). The percentage of nestin positive cells over the BLBP positive cells in the transection group was also higher than the normal group (the transection group:57.92±17.93%; the normal group: 23.26±9.85%; P< 0.01). And we observed more RGCs differentiated into MAP-2 positive neurons in the transection group (the transection group:46.13±14.92%; the normal group:29.13±10.07%; P< 0.01). Although there was no significant difference in the cell numbers between two groups when they differentiated into GFAP positive astrocytes and CNP positive oligodendrocyte, the cell bodies in the transection group were larger and the processes were also longer and thicker than the normal group.
Conclusions The results indicate that the denervated fimbria-fornix transected hippocampal extracts can significantly promote the proliferation of RGCs, enlarge the cell bodies, thicken and elongate the processes of BLBP positive RGCs which presenting the state of "activation" and containing the embryogenicity, and also can make RGCs present the identity of neural stem cells and differentiate into neurons, astrocytes and oligodendrocytes.
PartⅡTHE DIFFERENTION OF HIPPOCAMPAL RADIAL GLIAL CELLS INTO CHOLINERGIC NEURONS IN VITRO BY DENERVATED HIPPOCAMPAL EXTRACTS
Objective Fimbria-fornix transected hippocampal extracts were added into the cell culture medium to simulate the hippocampal internal environment in vitro, and to observe the situation of the RGCs differentiation into cholinergic neurons.
Methods Purified RGCs were divided into transection group and normal group. The DMEM/F12 medium supplemented with 5%(vol/vol) fimbria-fornix transected hippocampal extracts was added into the transection group, while the normal group the DMEM/F12 medium contains 5%(vol/vol) normal hippocampal extracts. BLBP/ChAT double-labeling immunofluorescence, Real-time PCR, and Western blot were used to observe the differentiation into cholinergic neurons, the levels of ChAT mRNA and protien in RGCs after 7 day inoculation.
Results Immunofluorescence assay of ChAT showed that the percentage of ChAT positive cholinergic neurons over the BLBP positive cells in the transection group(41.62±9.97%) was higher than the normal group(16.08±7.31%)(P< 0.01), and the cholinergic neuron bodies of transection group were larger and the processes were also longer and thicker than the normal group. The level of ChAT mRNA in the transection group was 5 times higher than the normal group (P< 0.01). The relative amount of ChAT protein in the transection group(0.1141±0.0380)was significant higher than that in the normal group(0.0423±0.0106)(P< 0.05), although both groups had low expression.
Conclusions The results indicate that the denervated hippocampal extracts after fimbria-fornix transection can significantly promote BLBP positive RGCs differentiate into cholinergic neurons.
引文
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