利用体细胞核移植技术生产转基因绒山羊的研究
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摘要
绒山羊体细胞核移植技术的研究目前处于刚刚起步阶段。本实验初步建立了转基因绒山羊克隆技术体系,并探讨了不同类型细胞对绒山羊克隆效率的影响,同时对转基因绒山羊进行了初步的鉴定以及生产性能检测。
1.绒山羊体细胞核移植技术的建立：
成功体外培养并且保存了四个胎儿来源的绒山羊胎儿成纤维细胞(caprine fetal fibroblast cells,CFC)以及两个成体耳尖成纤维细胞(caprine ear fibroblast cells, CEFC),并通过SRY-PCR方法鉴定了CFCs的性别。使用本实验室构建成功的真核表达载体通过脂质体转染CFC,经G418筛选获得了多个CFCs转基因细胞系(transgenic caprine fetal fibroblast cells, TCFC)。 TCFC高效表达外源标记基因红色荧光蛋白,且生长里旺盛,经PCR鉴定外源目的基因已经稳定整合入TCFC基因组中。
在检测转基因克隆胚胎发育能力时,绒山羊卵母细胞成熟率为72.6%(420/578),通过体细胞核移植获得重构胚胎419枚,卵裂率为86.1%(361/419),囊胚率为14.8%(62/419),克隆胚胎卵裂较为均匀并且高效表达红色荧光蛋白。2009年秋季进行了大规模的胚胎移植实验,共采集卵细胞6732枚,成熟率为65.1(4380/6732),融合处理重构胚胎3568枚,融合率为62.8%(2241/3568),卵裂率为54.9(1231/2241),移植胚胎924枚,移植受体273只(次),获得转基因及体细胞克隆动物20只。
2.供体细胞类型和受体羊条件对绒山羊体细胞核移植效率的影响：
本实验主要以不同种类成纤维细胞为核供体研究细胞类型以及受体山羊排卵数对绒山羊体细胞核移植效率的影响。分别使用不同个体来源胎儿成纤维细胞(Caprine Fetal Fibroblast Cells CFC)、耳尖成纤维细胞(Caprine Ear Fibroblast Cells, CEFC)、转基因胎儿成纤维细胞(Caprine Fetal Fibroblast Cells, TCFC)为核供体进行核移植操作,比较成纤维细胞来源、类型、转基因与否以及性别对融合率、卵裂率、出生率、出生后存活率以及死亡率的影响。共获得4943枚重构胚胎,其中融合3949枚,体外培养3737枚,共卵裂3094枚,1873枚形态较好的克隆胚胎进行了胚胎移植实验,共移植受体羊368只,出生48只,存活35只。不同类型与来源供体细胞其克隆效率不尽相同,其中胎儿成纤维细胞来源胚胎其克隆效率高于耳尖成纤维细胞来源胚胎,非转基因克隆胚胎效率高于转基因克隆胚胎,雄性细胞来源胚胎克隆效率高于雌性细胞来源胚胎,表明供体细胞的类型是决定克隆效率的重要因素之一。
克隆胚胎的卵裂率与羔羊出生率以及出生后存活率具有正相关性,并且早期克隆胚胎的发育能力与体细胞核移植效率具有一定的相关性。在分析不同年份克隆效率的波动时,发现受体母羊排卵点的多少也影响了克隆胚胎的移植受胎率。
3.转基因克隆绒山羊的鉴定与生产性能检测：
分别统计并分析了不同细胞来源的克隆羔羊妊娠天数与初生重的区别,并对转基因羔羊进行初步鉴定,检测分析了转基因克隆绒山羊的生产性能。结果表明不同细胞系来源的羔羊其妊娠天数也不尽相同,TCFC4细胞来源的克隆羔羊妊娠天数最长为平均157.5天,普通体细胞克隆羔羊妊娠天数最短只有152.3天。即使相同细胞系来源的克隆羔羊妊娠天数与出生重也不尽相同,尤其是在转基因细胞系来源的羔羊中存在较大的波动。
所有个体经PCR检测发现,外源目的基因IC片段与筛选基因Neo片段均稳定整合。最终获得的转基因动物的标记基因共出现三种表达情况：1、转基因羔羊的表型和耳尖分离培养的细胞均能观察到红色荧光；2、个体表型没有红色荧光,但分离培养的细胞具有荧光；3、个体表型和细胞均观察不到荧光。对同样来自CFC3细胞系的普通体细胞克隆羔羊、转IGF-1基因羔羊、转VEGF基因羔羊的生产性能进行了检测,发现转VEGF基因的绒山羊产绒量得到较大的提高。
It is just beganning of somatic cell nuclear transfer (SCNT) on cashmere goat, here we established the SCNT on transgenic cashmere goat, and the influence of cell types on cloning efficiency of cashmere goat was investigated. Finally the identification and production performance testing of cloned transgenic cashmere goats were proceeded.
1The establishment of cashmere goats SCNT:
The fibroblast cells from four caprine fetals(caprine fetal fibroblast cells,CFC) and2adults ears(caprine ear fibroblast cells, CEFC) were cultured, and the genders of them were distinguished by SRY-PCR. Transgenic caprine fetal fibroblast cells(transgenic caprine fetal fibroblast cells, TCFC) were obtained after transfected with eukaryotic expression vector and screened with G418. Red fluorescence could be detectable in the TCFCs, and the results of PCR demonstrated the exogous gene was integrated stably. SCNT was performed with these TCFCs utilized as the nucleus donors. The rate of oocyte IVM was72.6%(420/578), and419reconstructed embryos were harvested. In the piriod of in vitro development,86.1%(361/419) oocyte cleavage rate and14.8%(62/419) blastula rate were obtained, while the red fluorescence could be obseved in all blastulas. Another SCNT and embryo transfer were performed in2009. Here we harvested6732oocytes,65.1%(4380/6732) of them had matured in vitro,and3568enucleated oocytes were fusioned with TCFCs and CFCs, the rate of fusion and cleavage were62.8%(2241/3568) and54.9%(1231/2241) respectively. Totally924embryos were transfered into273recipients, and20kids were born.
2.The affect of donor cell types and recipients condition on cashmere goat SCNT efficiency:
The efficiency of donor cell types in cashmere goat SCNT was undefined. In this study, SCNT was performed using CFCs, CEFCs and TCFCs as donor cells to compare the influences of cell type, transgene and sex of nuclear donor fibroblast cells on the SCNT efficiency including fusion and cleavage rates of reconstructed embryos and the birth, postnatal survival and mortality rates of cloned kids. A total of4943reconstructed embryos were obtained. Among them,3949embryos were fused, and3737embryos were cultured in vitro leading to a total of3094cleavage embryos. Furthermore, embryo transplantation analysis was conducted on1873cloned embryos with relatively normal morphology, and368recipient goats were transplanted and48kids were born, of which,35kids survived. The SCNT efficiency of CFCs cell-derived embryos was higher than that of CEFCs-derived embryos, and the cloning efficiency of non-transgenic cloned embryo was higher than that of transgenic cloned embryos. Also, the SCNT efficiency of male cell-derived embryos was higher than that of female cell-derived embryos. High oocytes cleavage rates predicted high birth rates and postnatal survival rates. The results indicated that the donor cell types of cashmere goats affected the clone efficience, and connections existed between the clone embryos early development ability and the SCNT efficience. The physiology condition (ovulation number) of recipients also affected the clone efficience.
3. The Identification and production performance testing of cloned transgenic cashmere goats:
The days of gestation and weight of newborn of cloned kids from different cells were recorded and analyzed. We also identified the transgenic kids and tested the production preformance. There was difference between the kids from various cells on days of gestation:the ones cloned from RCFC4was the longest,157.5days, while the ones cloned from ordinary somatic cells was the shortest,152.3days. Even though the kids cloned with same donors were various in days of gestation and weight of newborn, and this was more obvious in ones from transgenic cells. The red fluorescence could be observed on phenotype of some kids and the cells separated from ear apex tissues of these ones, while it could be detected in cells rather than phenotype of some others, and no fluorescence could be found in both cells or phenotype of remains, although the the IC and Neo fragment included in the exogous gene could be amplified from all cells. In the production performance testing of goats cloned from ordinary, transfected with IGF-1and VEGF CFC3cells, we found that the VEGF transfected goats had the greater improvement on cashmere production.
1.绒山羊体细胞核移植技术的建立：
成功体外培养并且保存了四个胎儿来源的绒山羊胎儿成纤维细胞(caprine fetal fibroblast cells,CFC)以及两个成体耳尖成纤维细胞(caprine ear fibroblast cells, CEFC),并通过SRY-PCR方法鉴定了CFCs的性别。使用本实验室构建成功的真核表达载体通过脂质体转染CFC,经G418筛选获得了多个CFCs转基因细胞系(transgenic caprine fetal fibroblast cells, TCFC)。 TCFC高效表达外源标记基因红色荧光蛋白,且生长里旺盛,经PCR鉴定外源目的基因已经稳定整合入TCFC基因组中。
在检测转基因克隆胚胎发育能力时,绒山羊卵母细胞成熟率为72.6%(420/578),通过体细胞核移植获得重构胚胎419枚,卵裂率为86.1%(361/419),囊胚率为14.8%(62/419),克隆胚胎卵裂较为均匀并且高效表达红色荧光蛋白。2009年秋季进行了大规模的胚胎移植实验,共采集卵细胞6732枚,成熟率为65.1(4380/6732),融合处理重构胚胎3568枚,融合率为62.8%(2241/3568),卵裂率为54.9(1231/2241),移植胚胎924枚,移植受体273只(次),获得转基因及体细胞克隆动物20只。
2.供体细胞类型和受体羊条件对绒山羊体细胞核移植效率的影响：
本实验主要以不同种类成纤维细胞为核供体研究细胞类型以及受体山羊排卵数对绒山羊体细胞核移植效率的影响。分别使用不同个体来源胎儿成纤维细胞(Caprine Fetal Fibroblast Cells CFC)、耳尖成纤维细胞(Caprine Ear Fibroblast Cells, CEFC)、转基因胎儿成纤维细胞(Caprine Fetal Fibroblast Cells, TCFC)为核供体进行核移植操作,比较成纤维细胞来源、类型、转基因与否以及性别对融合率、卵裂率、出生率、出生后存活率以及死亡率的影响。共获得4943枚重构胚胎,其中融合3949枚,体外培养3737枚,共卵裂3094枚,1873枚形态较好的克隆胚胎进行了胚胎移植实验,共移植受体羊368只,出生48只,存活35只。不同类型与来源供体细胞其克隆效率不尽相同,其中胎儿成纤维细胞来源胚胎其克隆效率高于耳尖成纤维细胞来源胚胎,非转基因克隆胚胎效率高于转基因克隆胚胎,雄性细胞来源胚胎克隆效率高于雌性细胞来源胚胎,表明供体细胞的类型是决定克隆效率的重要因素之一。
克隆胚胎的卵裂率与羔羊出生率以及出生后存活率具有正相关性,并且早期克隆胚胎的发育能力与体细胞核移植效率具有一定的相关性。在分析不同年份克隆效率的波动时,发现受体母羊排卵点的多少也影响了克隆胚胎的移植受胎率。
3.转基因克隆绒山羊的鉴定与生产性能检测：
分别统计并分析了不同细胞来源的克隆羔羊妊娠天数与初生重的区别,并对转基因羔羊进行初步鉴定,检测分析了转基因克隆绒山羊的生产性能。结果表明不同细胞系来源的羔羊其妊娠天数也不尽相同,TCFC4细胞来源的克隆羔羊妊娠天数最长为平均157.5天,普通体细胞克隆羔羊妊娠天数最短只有152.3天。即使相同细胞系来源的克隆羔羊妊娠天数与出生重也不尽相同,尤其是在转基因细胞系来源的羔羊中存在较大的波动。
所有个体经PCR检测发现,外源目的基因IC片段与筛选基因Neo片段均稳定整合。最终获得的转基因动物的标记基因共出现三种表达情况：1、转基因羔羊的表型和耳尖分离培养的细胞均能观察到红色荧光；2、个体表型没有红色荧光,但分离培养的细胞具有荧光；3、个体表型和细胞均观察不到荧光。对同样来自CFC3细胞系的普通体细胞克隆羔羊、转IGF-1基因羔羊、转VEGF基因羔羊的生产性能进行了检测,发现转VEGF基因的绒山羊产绒量得到较大的提高。
It is just beganning of somatic cell nuclear transfer (SCNT) on cashmere goat, here we established the SCNT on transgenic cashmere goat, and the influence of cell types on cloning efficiency of cashmere goat was investigated. Finally the identification and production performance testing of cloned transgenic cashmere goats were proceeded.
1The establishment of cashmere goats SCNT:
The fibroblast cells from four caprine fetals(caprine fetal fibroblast cells,CFC) and2adults ears(caprine ear fibroblast cells, CEFC) were cultured, and the genders of them were distinguished by SRY-PCR. Transgenic caprine fetal fibroblast cells(transgenic caprine fetal fibroblast cells, TCFC) were obtained after transfected with eukaryotic expression vector and screened with G418. Red fluorescence could be detectable in the TCFCs, and the results of PCR demonstrated the exogous gene was integrated stably. SCNT was performed with these TCFCs utilized as the nucleus donors. The rate of oocyte IVM was72.6%(420/578), and419reconstructed embryos were harvested. In the piriod of in vitro development,86.1%(361/419) oocyte cleavage rate and14.8%(62/419) blastula rate were obtained, while the red fluorescence could be obseved in all blastulas. Another SCNT and embryo transfer were performed in2009. Here we harvested6732oocytes,65.1%(4380/6732) of them had matured in vitro,and3568enucleated oocytes were fusioned with TCFCs and CFCs, the rate of fusion and cleavage were62.8%(2241/3568) and54.9%(1231/2241) respectively. Totally924embryos were transfered into273recipients, and20kids were born.
2.The affect of donor cell types and recipients condition on cashmere goat SCNT efficiency:
The efficiency of donor cell types in cashmere goat SCNT was undefined. In this study, SCNT was performed using CFCs, CEFCs and TCFCs as donor cells to compare the influences of cell type, transgene and sex of nuclear donor fibroblast cells on the SCNT efficiency including fusion and cleavage rates of reconstructed embryos and the birth, postnatal survival and mortality rates of cloned kids. A total of4943reconstructed embryos were obtained. Among them,3949embryos were fused, and3737embryos were cultured in vitro leading to a total of3094cleavage embryos. Furthermore, embryo transplantation analysis was conducted on1873cloned embryos with relatively normal morphology, and368recipient goats were transplanted and48kids were born, of which,35kids survived. The SCNT efficiency of CFCs cell-derived embryos was higher than that of CEFCs-derived embryos, and the cloning efficiency of non-transgenic cloned embryo was higher than that of transgenic cloned embryos. Also, the SCNT efficiency of male cell-derived embryos was higher than that of female cell-derived embryos. High oocytes cleavage rates predicted high birth rates and postnatal survival rates. The results indicated that the donor cell types of cashmere goats affected the clone efficience, and connections existed between the clone embryos early development ability and the SCNT efficience. The physiology condition (ovulation number) of recipients also affected the clone efficience.
3. The Identification and production performance testing of cloned transgenic cashmere goats:
The days of gestation and weight of newborn of cloned kids from different cells were recorded and analyzed. We also identified the transgenic kids and tested the production preformance. There was difference between the kids from various cells on days of gestation:the ones cloned from RCFC4was the longest,157.5days, while the ones cloned from ordinary somatic cells was the shortest,152.3days. Even though the kids cloned with same donors were various in days of gestation and weight of newborn, and this was more obvious in ones from transgenic cells. The red fluorescence could be observed on phenotype of some kids and the cells separated from ear apex tissues of these ones, while it could be detected in cells rather than phenotype of some others, and no fluorescence could be found in both cells or phenotype of remains, although the the IC and Neo fragment included in the exogous gene could be amplified from all cells. In the production performance testing of goats cloned from ordinary, transfected with IGF-1and VEGF CFC3cells, we found that the VEGF transfected goats had the greater improvement on cashmere production.
引文
[1]Wilmut, I., A.E. Schnieke, J. Mc Whir, etc., Viable offspring derived from fetal and adult mammalian cells. Nature,1997.385(6619):810-3.
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[1]Wilmut, I., A.E. Schnieke, J. McWhir, etc., Viable offspring derived from fetal and adult mammalian cells. Nature,1997.385(6619):810-3.
[2]Schnieke, A.E., A.J. Kind, W.A. Ritchie, etc., Human factor IX transgenic sheep produced by transfer of nuclei from transfected fetal fibroblasts. Science, 1997.278(5346):2130-3.
[3]Lai, L., J.X. Kang, R. Li, etc., Generation of cloned transgenic pigs rich in omega-3 fatty acids. Nat Biotechnol,2006.24(4):435-6.
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[6]Kato, Y. and Y. Tsunoda, Role of the donor nuclei in cloning efficiency:can the ooplasm reprogram any nucleus? Int J Dev Biol,2010.54(11-12):1623-9.
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