鲁西黄牛胚胎骨骼肌卫星细胞分离培养及生物学特性研究
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摘要
骨骼肌卫星细胞（satellite cells，SCs）最早由Mauro在蛙胫前肌中发现，具有较强的自我更新、增殖和多向分化潜能，在体外培养条件下能跨胚层分化为多种细胞类型。鲁西黄牛是世界上著名的肉用牛，已列入国家级畜禽遗传资源重点保护品种名录。本研究旨在建立鲁西黄牛胚胎骨骼肌卫星细胞体外扩增培养体系，并探索其低、中、高代次细胞增殖、分化等生物学特性，结论如下：
（1）本研究选取鲁西黄牛四肢骨骼肌，采用机械分离法、联合酶消化法及差速贴壁法，成功分离获得牛SCs，优化了牛SCs体外扩增的最佳培养体系，并将牛SCs成功传至23代，冻存后牛SCs仍具有较高的细胞活率。
（2）免疫细胞化学染色检测牛SCs标志物表达，结果显示，P4、P9、P20代牛SCs均为Desmin、MyoD阳性表达；RT-PCR检测牛SCs标志物表达，结果显示P4、P9、P20代牛SCs均为Desmin、Pax7、c-Met、Myf5阳性表达，且灰度分析结果显示，不同代次SCs同一标志物表达量差异不显著，符合SCs标志物表达情况。
（3）P4、P9、P20代牛SCs生长曲线均呈标准“S”型，分别经历了潜伏期，对数生长期和平台期；不同代次SCs均具有较高的克隆形成能力，但克隆形成率随着细胞代次增加而降低；流式细胞仪分析SCs细胞周期，结果显示，随着细胞代次的增加，G0/G1期细胞比例增加，S期细胞比例下降。
（4）牛SCs染色体数目为2n=60，且染色体形态无畸变，除性染色体外，均为端着丝粒染色体，表明本试验分离获得的SCs均来自正常的鲁西黄牛胚胎个体。
（5）在不同的体外诱导条件下，诱导P4、P9、P20代牛SCs向成肌细胞、脂肪细胞、成骨细胞、神经细胞和表皮细胞分化。应用相应的染色方法检测诱导效果，结果显示，不同染色方法均呈阳性表达；RT-PCR检测SCs向不同细胞诱导后相应标志物的表达，包括成肌细胞标志物fast skeletal myosin，脂肪细胞标志物PPARγ、LPL，成骨细胞标志物osteopontin、Collage type Ⅰ，神经元标志物MAP2、nestin和表皮细胞标志物α6和CK19，结果显示，所有标志物均呈阳性表达。对目的基因表达量进行灰度分析发现，中、低代次SCs分化能力无显著差异，高代次SCs的分化能力显著降低。
实验证明，鲁西黄牛胚胎SCs适合在体外分离培养，经多次传代培养后，低、中、高代次SCs仍具有较强的自我更新、增殖和分化等生物学特性，且具有取材方便，获得细胞数量多等优点，可作为一种理想的种子细胞来保存鲁西黄牛这一珍贵的遗传资源，也为SCs在临床上的应用提供实验依据。
Muscle satellite cells (SCs) which provided with strong potential aspect ofself-renewal, proliferation and differentiation and could across the germinal layer todifferentiate into certain cell types in vitro were first discovered by Mauro in frog tibialisanterior muscle. Luxi cattle as a world known beef breed, has been enrolled into thecategories for key protected breed of national livestock and poultry genetic resources. Thepurpose of this study is to establish amplification system for Luxi cattle embryonic skeletalmuscle satellite cells in vitro, and to explore its biological characteristics such as cellproliferation, differentiation with low, medium and high three levels. The results are listedas follows:
(1) In this study, we isolated and purified bovine SCs in skeletal muscle from cattlelegs using mechanical separation, combined enzyme digestion and differential adhesionmethodologies. Then optimized best culture system for SCs amplification in vitro andsuccessfully passages to the23generation. It was founded that the SCs had higher cellviability after cryopreservation.
(2) The results of immunocytochemistry staining for detection of surface markersexpression in SCs showed that P4, P9, P20generations of Bovine SCs were positivelyexpressed with Desmin and MyoD. The results of RT-PCR detection for SCs surfacemarkers expression showed that all of P4, P9, P20generations of Bovine SCs werepositively expressed with Desmin, Pax7, c-Met, Myf5. And the result of Grayscale analysisshowed that the Variation of the same surface markers from different passages were notsignificant and coincidence with expressional situation of SCs surface marker.
(3) All kind of growth curves that went through certain stages including latency phase,logarithmic growth phase and stationary phase from P4, P9and P20. Generations of bovineSCs were presented typically “S” shape. Different passages of SCs all have a well colonyformation but the rates of colony formation were decreased gradually with the increasingnumber of cell passages. The results of flow cytometry analysis for SCs cell cycle showedthat the cell proportions were increased in G0/G1period and decreased in S periodaccording with the increased number of cell passages.
(4) The number of Bovine SCs chromosome is2n=60, and the morphology ofchromosomes were no distortion. The chromosomes are proximal to the centromerechromosomes except two sex chromosomes. Those evidence indicating that the isolated SCs in this expriment were all originated form normal Luxi cattle embryos.
(5) Induced P4, P9, P20generation of bovine SCs to differentiated into myobalsts,adipocyte, osteoblast, neurogenic and epidermal cells under different inducing conditions.Then applying corresponding staining methodology to detected the inducing affects. Theresults showed that the staining methods applied here were all expressed positively. AfterSCs were induced into different cells, we analyzed the expression of corresponding surfacemarkers including fast skeletal myosin in myotubules, PPARγ and LPL in adipocyte,osteopontin and collage type Ⅰin osteoblast, MAP2and nestin in neurogenic, and α6andCK19in epidermal cells by RT-PCR. Afterword, the results showed those markers were allexpressed positively. Grayscale analysis On expressional amount of target genes showedthat the differences of differential aspect in low-passage and middle-passage SCs were nosignificant difference. But the differential aspect of high-passage cells were reducedsignificantly.
These results confirmed that the Luxi cattle embryonic SCs were suitable for isolationand culture in vitro. The low, middle and high-passages SCs were still possess strongbiological characteristics such as self-renewal, proliferation and differentiation afterpassage cultured repeatedly. Besides, Cattle SCs had the advantage of easily and massiveobtained characteristics. So SCs would be used as an ideal seed cells type to protect thevaluable genetic resources Luxi cattle. In addition, it will provide experimental basis forthe clinical usage of SCs.
（1）本研究选取鲁西黄牛四肢骨骼肌，采用机械分离法、联合酶消化法及差速贴壁法，成功分离获得牛SCs，优化了牛SCs体外扩增的最佳培养体系，并将牛SCs成功传至23代，冻存后牛SCs仍具有较高的细胞活率。
（2）免疫细胞化学染色检测牛SCs标志物表达，结果显示，P4、P9、P20代牛SCs均为Desmin、MyoD阳性表达；RT-PCR检测牛SCs标志物表达，结果显示P4、P9、P20代牛SCs均为Desmin、Pax7、c-Met、Myf5阳性表达，且灰度分析结果显示，不同代次SCs同一标志物表达量差异不显著，符合SCs标志物表达情况。
（3）P4、P9、P20代牛SCs生长曲线均呈标准“S”型，分别经历了潜伏期，对数生长期和平台期；不同代次SCs均具有较高的克隆形成能力，但克隆形成率随着细胞代次增加而降低；流式细胞仪分析SCs细胞周期，结果显示，随着细胞代次的增加，G0/G1期细胞比例增加，S期细胞比例下降。
（4）牛SCs染色体数目为2n=60，且染色体形态无畸变，除性染色体外，均为端着丝粒染色体，表明本试验分离获得的SCs均来自正常的鲁西黄牛胚胎个体。
（5）在不同的体外诱导条件下，诱导P4、P9、P20代牛SCs向成肌细胞、脂肪细胞、成骨细胞、神经细胞和表皮细胞分化。应用相应的染色方法检测诱导效果，结果显示，不同染色方法均呈阳性表达；RT-PCR检测SCs向不同细胞诱导后相应标志物的表达，包括成肌细胞标志物fast skeletal myosin，脂肪细胞标志物PPARγ、LPL，成骨细胞标志物osteopontin、Collage type Ⅰ，神经元标志物MAP2、nestin和表皮细胞标志物α6和CK19，结果显示，所有标志物均呈阳性表达。对目的基因表达量进行灰度分析发现，中、低代次SCs分化能力无显著差异，高代次SCs的分化能力显著降低。
实验证明，鲁西黄牛胚胎SCs适合在体外分离培养，经多次传代培养后，低、中、高代次SCs仍具有较强的自我更新、增殖和分化等生物学特性，且具有取材方便，获得细胞数量多等优点，可作为一种理想的种子细胞来保存鲁西黄牛这一珍贵的遗传资源，也为SCs在临床上的应用提供实验依据。
Muscle satellite cells (SCs) which provided with strong potential aspect ofself-renewal, proliferation and differentiation and could across the germinal layer todifferentiate into certain cell types in vitro were first discovered by Mauro in frog tibialisanterior muscle. Luxi cattle as a world known beef breed, has been enrolled into thecategories for key protected breed of national livestock and poultry genetic resources. Thepurpose of this study is to establish amplification system for Luxi cattle embryonic skeletalmuscle satellite cells in vitro, and to explore its biological characteristics such as cellproliferation, differentiation with low, medium and high three levels. The results are listedas follows:
(1) In this study, we isolated and purified bovine SCs in skeletal muscle from cattlelegs using mechanical separation, combined enzyme digestion and differential adhesionmethodologies. Then optimized best culture system for SCs amplification in vitro andsuccessfully passages to the23generation. It was founded that the SCs had higher cellviability after cryopreservation.
(2) The results of immunocytochemistry staining for detection of surface markersexpression in SCs showed that P4, P9, P20generations of Bovine SCs were positivelyexpressed with Desmin and MyoD. The results of RT-PCR detection for SCs surfacemarkers expression showed that all of P4, P9, P20generations of Bovine SCs werepositively expressed with Desmin, Pax7, c-Met, Myf5. And the result of Grayscale analysisshowed that the Variation of the same surface markers from different passages were notsignificant and coincidence with expressional situation of SCs surface marker.
(3) All kind of growth curves that went through certain stages including latency phase,logarithmic growth phase and stationary phase from P4, P9and P20. Generations of bovineSCs were presented typically “S” shape. Different passages of SCs all have a well colonyformation but the rates of colony formation were decreased gradually with the increasingnumber of cell passages. The results of flow cytometry analysis for SCs cell cycle showedthat the cell proportions were increased in G0/G1period and decreased in S periodaccording with the increased number of cell passages.
(4) The number of Bovine SCs chromosome is2n=60, and the morphology ofchromosomes were no distortion. The chromosomes are proximal to the centromerechromosomes except two sex chromosomes. Those evidence indicating that the isolated SCs in this expriment were all originated form normal Luxi cattle embryos.
(5) Induced P4, P9, P20generation of bovine SCs to differentiated into myobalsts,adipocyte, osteoblast, neurogenic and epidermal cells under different inducing conditions.Then applying corresponding staining methodology to detected the inducing affects. Theresults showed that the staining methods applied here were all expressed positively. AfterSCs were induced into different cells, we analyzed the expression of corresponding surfacemarkers including fast skeletal myosin in myotubules, PPARγ and LPL in adipocyte,osteopontin and collage type Ⅰin osteoblast, MAP2and nestin in neurogenic, and α6andCK19in epidermal cells by RT-PCR. Afterword, the results showed those markers were allexpressed positively. Grayscale analysis On expressional amount of target genes showedthat the differences of differential aspect in low-passage and middle-passage SCs were nosignificant difference. But the differential aspect of high-passage cells were reducedsignificantly.
These results confirmed that the Luxi cattle embryonic SCs were suitable for isolationand culture in vitro. The low, middle and high-passages SCs were still possess strongbiological characteristics such as self-renewal, proliferation and differentiation afterpassage cultured repeatedly. Besides, Cattle SCs had the advantage of easily and massiveobtained characteristics. So SCs would be used as an ideal seed cells type to protect thevaluable genetic resources Luxi cattle. In addition, it will provide experimental basis forthe clinical usage of SCs.
引文
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