核质相互作用对供体核重编程的影响
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摘要
在农业生产和生物医学研究中,体细胞核移植都有着极其重要的应用,但是,克隆效率低下一直阻碍着体细胞核移植的进一步应用和发展。核移植的技术步骤较为复杂,很多相关的研究也一直围绕核移植操作流程中的受体卵母细胞的去核方法、融合过程、激活过程以及早期胚胎体外培养体系等方面进行了探讨。然而,由于克隆胚胎发育率的多变性和胎儿发育效率的低下性,除了从核移植的操作流程方面去探讨和改良外,同时还应该从生产重构胚胎的生物材料——即供体细胞和受体卵母细胞的来源、质量及表观遗传状态等——对于重编程的影响及二者之间的相互作用进行探讨和研究。
本研究首先优化了牛卵母细胞体外成熟培养体系。本实验对比了四种不同的激素配比对牛卵母细胞体外成熟的作用,结果显示添加人尿促性腺激素(HMG)的成熟实验结果稳定,17β-雌二醇(E2)对牛卵母细胞成熟有一定的促进作用,HMG+E2联合使用可以得到高效稳定的成熟结果;在此基础上,比较了表皮生长因子(EGF)对牛卵母细胞体外成熟和孤雌胚胎发育的影响,结果表明在成熟液中添加30 ng/ml EGF对牛卵母细胞的成熟质量、胚胎发育及降低胚胎细胞凋亡都有明显的促进作用。本研究最终确定的牛卵母细胞体外成熟培养液为:TCM-199中添加0.075 IU/ml HMG,1μg/ml E2,30 ng/ml EGF,10%(v/v)FBS。
接下来本研究对比了血清饥饿法与接触抑制法在调整供体细胞周期中对核移植效果产生的作用。结果发现,虽然两种方法在重构胚的融合率、卵裂率和囊胚发育率上并无太大的差异,二者均可以得到较好的结果;但是血清饥饿处理会引起胚胎凋亡率增加,导致核移植胚胎质量下降。细胞的传代会对核移植结果产生影响,以第2-5代细胞的效果最好,但当传代次数过多后则会造成重构胚发育能力的下降;也会引起核移植囊胚凋亡率上升。根据以上结果,本研究在后续实验中均使用接触抑制法处理的第2-5代细胞为供体细胞。
随后,本研究分别应用未成熟卵母细胞抽提物、成熟卵母细胞抽提物和孤雌激活的卵母细胞抽提物预处理供体细胞,通过对比处理细胞的全能性基因表达和脱乙酰化程度检测三种胞质抽提物对供体细胞的重编程效果。随后进行核移植实验,以正常核移植胚胎为对照比较了来自于三组不同抽提物预处理供体细胞的重构胚的体外发育情况和囊胚质量,从而分析牛卵胞质抽提物对供体细胞的重编程作用对体细胞核移植的影响。结果如下,三组胞质抽提物都能有效的重编程供体细胞;在三组抽提物中,成熟卵胞质抽提物使得供体核的表观遗传状态与受体胞质更为接近,因此对克隆胚胎发育率和囊胚质量有显著的促进作用。这些结果表明,用牛卵胞质抽提物预处理供体细胞能提高重构胚体外发育能力,相对于将供体核重编程到一个更低的分化状态,将供体核重编程到一个与受体胞质相一致的状态更有利于克隆胚胎的发育。
本研究设计使用小鼠和牛两种在基因表达调控和发育程序上有着巨大差异的物种研究早期胚胎中供体核与胞质在指导发育上的作用,构建了小鼠-牛异种克隆胚胎,检测了异种胚胎的体外发育时间、胚胎形态、合子激活基因的表达情况,结果表明异种克隆胚胎在合子基因激活(ZGA)前的发育是由牛的卵胞质控制的,在发育时序和胚胎形态上都表现出牛胚胎的特性,同时小鼠的合子基因激活也受到牛卵胞质的作用而在8-细胞期才得以表达,并且激活是不完全的。小鼠的合子基因激活受到牛卵胞质的影响而延迟和不完全激活是造成小鼠-牛异种克隆胚阻滞在8-细胞期的一个原因。
最后,本研究设计使用38.5℃正常培养的孤雌激活胚胎与41℃培养7 h的热激孤雌激活胚胎进行核质置换,分别构建热激核-正常胞质重构胚以及热激胞质-正常核重构胚,检测了核质置换胚胎的体外发育率、合子激活基因的表达和胚胎凋亡情况,以研究早期胚胎核与胞质对热激的应激反应,以及产生应激后二者之间的相互作用。结果如下,当正常核移入热激的卵胞质,重构胚的发育率显著下降;热激后的核质置换胚胎中存在合子基因激活不完全的现象,这种情况主要存在于热激胞质-正常核重构胚中;热激能诱导胚胎发生细胞凋亡,早期胚胎的胞质对热激更敏感并在后期的诱导凋亡中起到主导作用。这些结果表明,虽然热激对核和胞质都有损伤,但胞质对热激损伤更为敏感。早期胚胎受到热激后导致发育能力降低的易感性主要是由于胞质所引起的;早期胚胎受到热激可能会损伤到卵胞质中的母源因子,即使与正常核构成重构胚也依然会导致合子基因表达异常;并且胞质受到热激损伤会在热激诱导的凋亡机制中起到比核更大的作用。
Somatic cell nuclear transfer (SCNT) has wide applications in agriculture and biomedical research. Yet low efficiency of cloning remains the major obstacle to the development and widespread use of this technology. SCNT includes complicated protocols; therefore many variables will have great effects on nuclear transfer efficiency. Intensive investigation has been performed on the nuclear transfer procedures, which includes but is not limited to enucleation, fusion, activation, and in vitro culture system of embryos. However, in vitro development rates of cloned embryos vary extensively and cloned fetus often fails to develop to term. Besides improvement of the SCNT procedures, thus, attention should be paid to the source, quality and epigenetic state of initial biological materials for producing cloned embryos. Therefore, investigation on donor cells, oocytes and nuclear-cytoplasmic interaction was performed in the present study.
In the present study, we first optimized the in vitro maturation system of bovine oocytes. Bovine oocytes were exposed to four different hormonal regimes in TCM-199 medium. No significant difference was observed in maturation rates in four groups. Human menopausal gonadotropin (HMG) provided steady maturation results in replicates. Maturation of oocytes was promoted by supplementation with 17β-estradiol to some extent. Combination of HMG and E2 gave rise to steady and efficient mature results. Then we studied the effect of epidermal growth factor (EGF) present in the above defined medium on bovine oocyte maturation, in vitro development and quality of parthenogenetic embryos. The results showed that the presence of EGF at 30 ng/ml concentration significantly increased maturation rate and blastocyst rate and reduced apoptotic cells in parthenogenetic blastocysts. Based on the above results, the final oocyte maturation (OM) solution were TCM-199 supplemented with 0.075 IU/ml HMG,1μg/ml E2,30 ng/ml EGF,10%(v/v)FBS.
Then the present work investigated changes to G0/G1 phase of donor cells treated with serum starvation or contact inhibition on in vitro development and quality of nuclear transfer embryos. No significant difference was observed in fused oocyte rate, cleavage rate and blastocyst rate between serum starvation and contact inhibition. However, more apoptotic cells were detected in blastocysts derived from donor cells treated with serum starvation. The passages of donor cells had an adverse effect on nuclear transfer. The 2nd to 5th passages of cell line were better than the other passages. Especially, repetitious passage impaired embryo competence and increased apoptotic rate in nuclear transfer blastocysts. Therefore, in this study, the 2nd to 5th passages of cell line were utilized as nuclei donors.
The present study investigated the effects of donor cells pretreated with oocyte extracts on in vitro development of cloned embryos. Bovine fibroblasts were exposed to immature, mature and parthenogenetic oocyte extracts respectively before nuclear transfer. The detectable expression of totipotent genes and global deacetylation in the treated cells showed that extracts could reprogram fibroblasts. Although all three groups of extracts exhibited reprogramming capacity, embryo development was not compliant with reprogramming effect. Improved quality and development of blastocysts were observed only in the mature extract treated group. We demonstrated that pretreatment of donor cells with mature oocyte extract improved in vitro development of cloned embryos. The present study demonstrated that it was synchronization between the epigenetic state of donor nuclei and recipient cytoplasm that played a more important role in nuclear-cytoplasmic interaction. Our results suggested that, rather than reprogramming donor nuclei to a lower differentiation state, reprogramming donor nuclei to a state synchronised with recipient cytoplasm before nuclear transfer would be beneficial for the development of cloned embryos.
The present study utilized mouse and bovine, huge differences in gene expression patterns and developmental program existed between the two species, to investigate the modulation of donor nuclei and cytoplasm on early embryogenesis. Mouse-bovine interspecies nuclear transfer embryos were constructed. The timing of in vitro development and morphology of heterogenous embryos and expression patterns of zygotic activated genes in heterogenous embryos were assessed. The results showed that the development of interspecies cloned embryos was modulated by bovine ooplasm before zygotic gene activation (ZGA). At this time, heterogenous embryos presented bovine phenotype in the timing and morphology. ZGA of mouse nuclei were controlled by bovine ooplasm and would be activated until 8-cell embryo, furthermore, this activation was incompletely. Retardant and incomplete activation of zygotic genes on mouse nuclei might result in 8-cell block of mouse-bovine interspecies nuclear transfer embryos.
Finally, the present study investigated the differential detrimental effects of heat shock on the nuclear or cytoplasmic component of bovine embryos during the first cell cycle. The parthenogenetic embryos were incubated at 38.5 or 41℃for 7 h, respectively; then the nuclei of embryos cultured at 41℃were transferred into enucleated 38.5℃cytoplasts to construct heat shock nuclei-normal cytoplasm embryos (HN embryos) and 38.5℃nuclei were transferred into 41℃cytoplasts to construct heat shock cytoplasm-normal nuclei embryos (HC embryos). In vitro culture showed that blastocyst formation rate for HC embryos decreased significantly compared to control parthenogenetic embryos and HN embryos. Incomplete activation of zygotic genes was observed in nuclear exchange embryos, especially in HC embryos. It suggested that heat shock might disrupt some cytoplasmic factors required for zygotic gene activation. Heat shock induced apoptosis was observed in both HN and HC blastocysts and the incidence of apoptosis in HC blastocysts was significantly higher than that in HN embryos. These results suggested that, although heat shock injuries existed in both nuclear and cytoplasmic components; cytoplasmic component is more susceptible to heat shock than nuclear component.
本研究首先优化了牛卵母细胞体外成熟培养体系。本实验对比了四种不同的激素配比对牛卵母细胞体外成熟的作用,结果显示添加人尿促性腺激素(HMG)的成熟实验结果稳定,17β-雌二醇(E2)对牛卵母细胞成熟有一定的促进作用,HMG+E2联合使用可以得到高效稳定的成熟结果;在此基础上,比较了表皮生长因子(EGF)对牛卵母细胞体外成熟和孤雌胚胎发育的影响,结果表明在成熟液中添加30 ng/ml EGF对牛卵母细胞的成熟质量、胚胎发育及降低胚胎细胞凋亡都有明显的促进作用。本研究最终确定的牛卵母细胞体外成熟培养液为:TCM-199中添加0.075 IU/ml HMG,1μg/ml E2,30 ng/ml EGF,10%(v/v)FBS。
接下来本研究对比了血清饥饿法与接触抑制法在调整供体细胞周期中对核移植效果产生的作用。结果发现,虽然两种方法在重构胚的融合率、卵裂率和囊胚发育率上并无太大的差异,二者均可以得到较好的结果;但是血清饥饿处理会引起胚胎凋亡率增加,导致核移植胚胎质量下降。细胞的传代会对核移植结果产生影响,以第2-5代细胞的效果最好,但当传代次数过多后则会造成重构胚发育能力的下降;也会引起核移植囊胚凋亡率上升。根据以上结果,本研究在后续实验中均使用接触抑制法处理的第2-5代细胞为供体细胞。
随后,本研究分别应用未成熟卵母细胞抽提物、成熟卵母细胞抽提物和孤雌激活的卵母细胞抽提物预处理供体细胞,通过对比处理细胞的全能性基因表达和脱乙酰化程度检测三种胞质抽提物对供体细胞的重编程效果。随后进行核移植实验,以正常核移植胚胎为对照比较了来自于三组不同抽提物预处理供体细胞的重构胚的体外发育情况和囊胚质量,从而分析牛卵胞质抽提物对供体细胞的重编程作用对体细胞核移植的影响。结果如下,三组胞质抽提物都能有效的重编程供体细胞;在三组抽提物中,成熟卵胞质抽提物使得供体核的表观遗传状态与受体胞质更为接近,因此对克隆胚胎发育率和囊胚质量有显著的促进作用。这些结果表明,用牛卵胞质抽提物预处理供体细胞能提高重构胚体外发育能力,相对于将供体核重编程到一个更低的分化状态,将供体核重编程到一个与受体胞质相一致的状态更有利于克隆胚胎的发育。
本研究设计使用小鼠和牛两种在基因表达调控和发育程序上有着巨大差异的物种研究早期胚胎中供体核与胞质在指导发育上的作用,构建了小鼠-牛异种克隆胚胎,检测了异种胚胎的体外发育时间、胚胎形态、合子激活基因的表达情况,结果表明异种克隆胚胎在合子基因激活(ZGA)前的发育是由牛的卵胞质控制的,在发育时序和胚胎形态上都表现出牛胚胎的特性,同时小鼠的合子基因激活也受到牛卵胞质的作用而在8-细胞期才得以表达,并且激活是不完全的。小鼠的合子基因激活受到牛卵胞质的影响而延迟和不完全激活是造成小鼠-牛异种克隆胚阻滞在8-细胞期的一个原因。
最后,本研究设计使用38.5℃正常培养的孤雌激活胚胎与41℃培养7 h的热激孤雌激活胚胎进行核质置换,分别构建热激核-正常胞质重构胚以及热激胞质-正常核重构胚,检测了核质置换胚胎的体外发育率、合子激活基因的表达和胚胎凋亡情况,以研究早期胚胎核与胞质对热激的应激反应,以及产生应激后二者之间的相互作用。结果如下,当正常核移入热激的卵胞质,重构胚的发育率显著下降;热激后的核质置换胚胎中存在合子基因激活不完全的现象,这种情况主要存在于热激胞质-正常核重构胚中;热激能诱导胚胎发生细胞凋亡,早期胚胎的胞质对热激更敏感并在后期的诱导凋亡中起到主导作用。这些结果表明,虽然热激对核和胞质都有损伤,但胞质对热激损伤更为敏感。早期胚胎受到热激后导致发育能力降低的易感性主要是由于胞质所引起的;早期胚胎受到热激可能会损伤到卵胞质中的母源因子,即使与正常核构成重构胚也依然会导致合子基因表达异常;并且胞质受到热激损伤会在热激诱导的凋亡机制中起到比核更大的作用。
Somatic cell nuclear transfer (SCNT) has wide applications in agriculture and biomedical research. Yet low efficiency of cloning remains the major obstacle to the development and widespread use of this technology. SCNT includes complicated protocols; therefore many variables will have great effects on nuclear transfer efficiency. Intensive investigation has been performed on the nuclear transfer procedures, which includes but is not limited to enucleation, fusion, activation, and in vitro culture system of embryos. However, in vitro development rates of cloned embryos vary extensively and cloned fetus often fails to develop to term. Besides improvement of the SCNT procedures, thus, attention should be paid to the source, quality and epigenetic state of initial biological materials for producing cloned embryos. Therefore, investigation on donor cells, oocytes and nuclear-cytoplasmic interaction was performed in the present study.
In the present study, we first optimized the in vitro maturation system of bovine oocytes. Bovine oocytes were exposed to four different hormonal regimes in TCM-199 medium. No significant difference was observed in maturation rates in four groups. Human menopausal gonadotropin (HMG) provided steady maturation results in replicates. Maturation of oocytes was promoted by supplementation with 17β-estradiol to some extent. Combination of HMG and E2 gave rise to steady and efficient mature results. Then we studied the effect of epidermal growth factor (EGF) present in the above defined medium on bovine oocyte maturation, in vitro development and quality of parthenogenetic embryos. The results showed that the presence of EGF at 30 ng/ml concentration significantly increased maturation rate and blastocyst rate and reduced apoptotic cells in parthenogenetic blastocysts. Based on the above results, the final oocyte maturation (OM) solution were TCM-199 supplemented with 0.075 IU/ml HMG,1μg/ml E2,30 ng/ml EGF,10%(v/v)FBS.
Then the present work investigated changes to G0/G1 phase of donor cells treated with serum starvation or contact inhibition on in vitro development and quality of nuclear transfer embryos. No significant difference was observed in fused oocyte rate, cleavage rate and blastocyst rate between serum starvation and contact inhibition. However, more apoptotic cells were detected in blastocysts derived from donor cells treated with serum starvation. The passages of donor cells had an adverse effect on nuclear transfer. The 2nd to 5th passages of cell line were better than the other passages. Especially, repetitious passage impaired embryo competence and increased apoptotic rate in nuclear transfer blastocysts. Therefore, in this study, the 2nd to 5th passages of cell line were utilized as nuclei donors.
The present study investigated the effects of donor cells pretreated with oocyte extracts on in vitro development of cloned embryos. Bovine fibroblasts were exposed to immature, mature and parthenogenetic oocyte extracts respectively before nuclear transfer. The detectable expression of totipotent genes and global deacetylation in the treated cells showed that extracts could reprogram fibroblasts. Although all three groups of extracts exhibited reprogramming capacity, embryo development was not compliant with reprogramming effect. Improved quality and development of blastocysts were observed only in the mature extract treated group. We demonstrated that pretreatment of donor cells with mature oocyte extract improved in vitro development of cloned embryos. The present study demonstrated that it was synchronization between the epigenetic state of donor nuclei and recipient cytoplasm that played a more important role in nuclear-cytoplasmic interaction. Our results suggested that, rather than reprogramming donor nuclei to a lower differentiation state, reprogramming donor nuclei to a state synchronised with recipient cytoplasm before nuclear transfer would be beneficial for the development of cloned embryos.
The present study utilized mouse and bovine, huge differences in gene expression patterns and developmental program existed between the two species, to investigate the modulation of donor nuclei and cytoplasm on early embryogenesis. Mouse-bovine interspecies nuclear transfer embryos were constructed. The timing of in vitro development and morphology of heterogenous embryos and expression patterns of zygotic activated genes in heterogenous embryos were assessed. The results showed that the development of interspecies cloned embryos was modulated by bovine ooplasm before zygotic gene activation (ZGA). At this time, heterogenous embryos presented bovine phenotype in the timing and morphology. ZGA of mouse nuclei were controlled by bovine ooplasm and would be activated until 8-cell embryo, furthermore, this activation was incompletely. Retardant and incomplete activation of zygotic genes on mouse nuclei might result in 8-cell block of mouse-bovine interspecies nuclear transfer embryos.
Finally, the present study investigated the differential detrimental effects of heat shock on the nuclear or cytoplasmic component of bovine embryos during the first cell cycle. The parthenogenetic embryos were incubated at 38.5 or 41℃for 7 h, respectively; then the nuclei of embryos cultured at 41℃were transferred into enucleated 38.5℃cytoplasts to construct heat shock nuclei-normal cytoplasm embryos (HN embryos) and 38.5℃nuclei were transferred into 41℃cytoplasts to construct heat shock cytoplasm-normal nuclei embryos (HC embryos). In vitro culture showed that blastocyst formation rate for HC embryos decreased significantly compared to control parthenogenetic embryos and HN embryos. Incomplete activation of zygotic genes was observed in nuclear exchange embryos, especially in HC embryos. It suggested that heat shock might disrupt some cytoplasmic factors required for zygotic gene activation. Heat shock induced apoptosis was observed in both HN and HC blastocysts and the incidence of apoptosis in HC blastocysts was significantly higher than that in HN embryos. These results suggested that, although heat shock injuries existed in both nuclear and cytoplasmic components; cytoplasmic component is more susceptible to heat shock than nuclear component.
引文
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