去乙酰化抑制剂对五指山小型猪体细胞核移植胚胎发育能力的影响
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摘要
目前通过体细胞核移植(SCNT)技术已经成功获得多种哺乳动物的克隆后代,但动物克隆的成功率仍较低,大量的研究表明其可能是SCNT后表观遗传重编程的失败所致。组蛋白的高乙酰化可以增强转录因子与核小体之间的相互作用,从而有利于基因转录的启动。组蛋白去乙酰化酶抑制剂制(HDACi),丙戊酸(VPA)、 Scriptaid等能增加基因组中组蛋白的乙酰化水平。本研究利用VPA、Scriptaid能够抑制重构胚早期组蛋白去乙酸化的作用这一机制,对体细胞核移植后的重构胚进行不同浓度及不同时间的处理,观察并探讨其对克隆胚胎体内及体外发育能力的影响。
第一章：本研究以五指山小型猪成纤维细胞为供体细胞进行SCNT重构胚胎的构建,重构胚胎激活后培养在添加不同浓度(0～8nM) VPA的胚胎培养液中进行不同时间(0-48h)的体外培养,探讨了HDACi-VPA对五指山小型猪SCNT克隆胚胎的体外及体内发育能力方面的影响。实验1研究不同浓度的VPA在处理核移植胚胎24h条件下对其克隆胚胎体外发育力的影响。实验结果表明2mM VPA处理组(21.5%)克隆胚胎的囊胚发育率显著高于对照组、4mM处理组及8mM处理组(21.5vs10.5,12.6,17.2, p<0.05)。实验2选择实验1中筛选出的最佳浓度2mM为本实验的处理浓度,研究此浓度在不同添加处理时间的条件下对化学激活后核移植克隆胚胎的体外发育能力的影响。实验结果表明VPA处理24h组(20.7%)的囊胚发育率显著高于其它各组(对照组9.2%,12h处理组12.1%,48h处理组9.1%,p<0.05)。实验3将1～8细胞期的2mM VPA处理24h克隆胚胎(192～216枚)和未经处理(对照组)的克隆胚胎(179～225枚)分别移植到7头自然发情的受体母猪中。经B超检测其中VPA处理的3头受体母猪全部妊娠,2头足月分娩共产11头克隆仔猪,对照组的4头受体母猪3头妊娠,2头足月分娩共产12头克隆仔猪。实验发现VPA处理组克隆仔猪初生重及出生后3个月生存率,均低于未处理组克隆仔猪(675.2±185.3)。综合以上结果表明,VPA处理体细胞核移植胚胎可以提高克隆胚胎的体外囊胚率,对于体内发育能力有必要进一步研究。
第二章：本研究探讨了HDACi-SCR对五指山小型猪SCNT克隆胚胎的体外及体内发育能力方面的影响。实验1研究不同浓度的SCR在处理核移植胚胎24h条件下对其克隆胚胎体外发育力的影响。实验结果表明处理250nM处理组(20.0%)较对照组(12.2%)、50nM处理组(8.9%)及500nM处理组(11.9%)显著提高,p<0.05。实验2研究250nM SCR在不同的处理时间的条件下对化学激活后克隆胚胎的外发育能力的影响。实验结果发现SCR处理24h组(22.2%)克隆胚胎的囊胚发育率较12h处理组(10.0%)显著提高,p<0.05,与48h处理组(18.5%)没有差异,p>0.05。实验3分别以SCR、VPA及SCR+VPA处理化学激活后的体细胞核移植克隆胚胎,研究并探讨了SCR和VPA联合处理对克隆胚胎的体外发育力的影响。结果表明250nM SCR和2mM VPA联合处理24h组与对照组相比克隆胚胎的囊胚率显著提高(18.3vs9.2%,p<0.05),但与250nM SCR处理24h组(19.9%)及2mM VPA处理24h组(18.9%)比较囊胚率无显著性差异(p>0.05)。实验4研究了SCR对体细胞核移植克隆胚胎的体内发育情况的影响,将250nM SCR处理24h与对照组经过体细胞核移植后获得克隆胚胎分别移植于代孕母后,对照组移植的受体可成功产下健康的克隆仔猪,而SCR处理组并未获得克隆仔猪。综合以上结果表明,SCR处理体细胞核移植胚胎可以提高克隆胚胎的体外囊胚率,对于体内发育能力有必要进一步研究。此外,SCR与VPA联合处理对体细胞核移植克隆胚胎的体外发育无协同作用。
第三章：增强型绿色荧光蛋白(EGFP)在转基因动物的研究中可用来标记目的基因、筛选阳性胚胎、建立荧光动物模型：如将EGFP或红色荧光蛋白基因插入到猪的基因组内,可以很方便的对细胞和组织内某些因子的表达和代谢进行跟踪,从而为基础性研究提供便利。本试验以获得的转EGFP基因阳性细胞作为供体细胞,经核移植和电融合后在2mM VPA处理24h的转基因重构胚移入去核的卵母细胞中,成功地获得了转基因囊胚,并比较克隆胚胎体外及体内发育情况。结果表明GFP转基因重构胚重构胚体外囊胚率为15.1%,与未转基因组11.4%相比无显差异著(P>0.05)。将1～8细胞阶段重构胚胎移植到3头自然发情的五指山小型猪输卵管内,2头未发育到期而返情,1头怀孕106天后,成功产下4头克隆猪。其中2头为死胎,体重分别为247g和313g,其余2头在分娩后由于饲养管理不善死亡,体重分别为366g和573g。经过对猪皮肤组织外来基因检测及受体母猪、克隆猪与供体细胞的13个多态性位点的微卫星检测结果表明,其中2头具有绿色荧光遗传特征,4头克隆猪个体与供核细胞具有完全相同的多态性,而克隆猪个体与代孕母猪的多态性不同,证明克隆仔猪与代孕母猪无亲缘关系。研究结果为转基因五指山小型猪体细胞克隆的技术体系建立提供参考,从而为人类疾病发病机理研究以及生产人类遗传疾病的基因模型动物奠定基础。
Epigenetic modification influences reprogramming and subsequent development of somatic cell nuclear transfer(SCNT)embryos. Such modification includes an increase in histone acetylation. Histone deacetylase inhibitors(HDACi), such as valproic acid (VPA) and scriptaid (SCR), have been known to maintain a high cellular level of histone acetylation. Hence, treatment of nuclear transfer embryos with HDACi may increase the efficiency of cloning. The present study attempted to examine the effects of VPA and SCR with regard to the potency of enhancement of in vitro and in vivo development of Wuzhishan miniature pig SCNT embryos.
In CHAPTER1. The aim of the present study was to examine the effects of VPA, a HDACi on in vitro and in vivo development of Wuzhishan miniature pig SCNT embryos. Experiment1compared the in vitro developmental competence of nuclear transfer embryos treated with various concentrations of VPA for24h. Embryos treated with2mM VPA for24h reported a significantly increased rate of blastocyst formation compared with controls or4mM VPA and8mM VPA treated groups (21.5vs10.5,12.6and17.2%). Experiment2examined the in vitro developmental competence of nuclear transfer embryos treated with2mM VPA for different time periods following chemical activation. Embryos treated for24h reported higher rates of blastocyst formation than the controls or treated for4h and48h groups (20.7vs9.2,12.1and9.1%). In experiment3, an average of207(192to216) nuclear transfer embryos from the VPA-treated group were transferred to surrogate mothers, resulting in three pregnancies. Two of the surrogates delivered a total of11live piglets, the nine of11piglets in the VPA-treated group died in an unknown reason within1day to5days after birch. Untreated control embryos (average,205(179to225)) transferred to four surrogate mothers resulted in three pregnancies, two of which delivered a total of12live offspring, four of the12piglets in the VPA-untreated group died in an unknown reason within1day to3days, eight of12of the12piglets in the VPA-untreated group have survived over3or4months. The average birth weight of the two litters from the VPA-treated group was tended to be lower than that from the control groups (551.6g vs675.2g). These results suggest that VPA treatment increases the rate of blastocyst formation for SCNT embryos, both the VPA-treated and the untreated clones can develop to term in pigs, but the offspring from VPA-treated embryos survived to adulthood lower than the offspring from VPA-treated embryos(18.2vs66.9%; P<0.05).
In CHAPTER2. The aim of the present study was to examine the effects of SCR, a HDACi, on the in vitro and in vivo development of Wuzhishan miniature pig SCNT embryos. Experiment1compared the in vitro developmental competence of nuclear transfer embryos treated with various concentrations of SCR for24h. Embryos treated with250nM SCR showed a significantly increased rate of blastocyst formation compared with controls or embryos treated with50nM or500nM SCR (20.0vs12.2,8.9, and11.9%). Experiment2examined the in vitro developmental competence of nuclear transfer embryos treated with250nM SCR for different time periods following chemical activation. Embryos treated for24h showed higher rates of blastocyst formation than controls or embryos treated for12h or48h (22.2vs11.5,10.0, and18.5%). Experiment3directly compared the effects of SCR and VPA treatment and examined the additive effect of SCR and VPA on nuclear transfer embryos following chemical activation. Embryos treated with250nM SCR or2mM VPA for24h showed a significantly increased rate of blastocyst formation compared with controls (19.9and18.3vs9.2%), but not increased with a combination of250nM SCR and2mM VPA for24h (18.9vs19.9,18.3%). In experiment4, nuclear transfer embryos treated with250nM SCR for24h and non-treated controls were transferred to surrogate mothers, resulting in the birth of healthy cloned pups from the controls but no births in the SCR-treated group. These results suggest that SCR treatment increases the rate of blastocyst formation in somatic cell nuclear transfer embryos and affects their subsequent growth. Also, co-treatment with VPA and SCR did not improve the blastocyst formation rate.
In CHAPTER3. Genetically engineered pigs with cell markers such as fluorescent proteins are highly useful in lines of research that include the tracking of transplanted cells or tissues. It has been demonstrated that increased histone acetylation in SCNT embryos, by applying a histone deacetylase(HDAC) inhibitor such as VPA significantly enhances the developmental competence in our previous research. Herein, we report that the re constructed embryos treatment with2mM VPA for24h, and compared the development of SCNT embryos to the blastocyst stage when Wuzhishan miniature pig ear fibroblasts as donor cells, there have no significant differences were observed in the cleavage rates and blastocyste formation rates in non-trasfected or transfected ear fibroblasts (72.6,15.1and74.5,11.4%). we used the ear fibroblasts of miniature pigs transfected with EGFP as the donor cells, and transplanted reconstructed embryos (at the1to8cell stage) into the oviducts of three sexually mature Wuzhishan miniature pigs that were naturally estrus. One recipient successfully birthed two stillborn and two live piglets after116days of pregnancy. We investigated the presence of the foreign gene in the skin tissues of the piglets and tested13microsatellite polymorphisms in the recipient pigs, cloned offspring and donor cells. The results indicated that two cloned offspring expressed EGFP and all four cloned offspring had the same polymorphisms at all13microsatellites as the donor cell. However, the microsatellites varied between the cloned offspring and the recipient pigs, indicating the absence of a blood relationship between the two. The study therefore establishes a system to obtain transgenic Wuzhishan inbred miniature pigs by SCNT, thereby providing a basis to study the pathologic mechanisms of human disease and producing animal models for human genetic disorders.
第一章：本研究以五指山小型猪成纤维细胞为供体细胞进行SCNT重构胚胎的构建,重构胚胎激活后培养在添加不同浓度(0～8nM) VPA的胚胎培养液中进行不同时间(0-48h)的体外培养,探讨了HDACi-VPA对五指山小型猪SCNT克隆胚胎的体外及体内发育能力方面的影响。实验1研究不同浓度的VPA在处理核移植胚胎24h条件下对其克隆胚胎体外发育力的影响。实验结果表明2mM VPA处理组(21.5%)克隆胚胎的囊胚发育率显著高于对照组、4mM处理组及8mM处理组(21.5vs10.5,12.6,17.2, p<0.05)。实验2选择实验1中筛选出的最佳浓度2mM为本实验的处理浓度,研究此浓度在不同添加处理时间的条件下对化学激活后核移植克隆胚胎的体外发育能力的影响。实验结果表明VPA处理24h组(20.7%)的囊胚发育率显著高于其它各组(对照组9.2%,12h处理组12.1%,48h处理组9.1%,p<0.05)。实验3将1～8细胞期的2mM VPA处理24h克隆胚胎(192～216枚)和未经处理(对照组)的克隆胚胎(179～225枚)分别移植到7头自然发情的受体母猪中。经B超检测其中VPA处理的3头受体母猪全部妊娠,2头足月分娩共产11头克隆仔猪,对照组的4头受体母猪3头妊娠,2头足月分娩共产12头克隆仔猪。实验发现VPA处理组克隆仔猪初生重及出生后3个月生存率,均低于未处理组克隆仔猪(675.2±185.3)。综合以上结果表明,VPA处理体细胞核移植胚胎可以提高克隆胚胎的体外囊胚率,对于体内发育能力有必要进一步研究。
第二章：本研究探讨了HDACi-SCR对五指山小型猪SCNT克隆胚胎的体外及体内发育能力方面的影响。实验1研究不同浓度的SCR在处理核移植胚胎24h条件下对其克隆胚胎体外发育力的影响。实验结果表明处理250nM处理组(20.0%)较对照组(12.2%)、50nM处理组(8.9%)及500nM处理组(11.9%)显著提高,p<0.05。实验2研究250nM SCR在不同的处理时间的条件下对化学激活后克隆胚胎的外发育能力的影响。实验结果发现SCR处理24h组(22.2%)克隆胚胎的囊胚发育率较12h处理组(10.0%)显著提高,p<0.05,与48h处理组(18.5%)没有差异,p>0.05。实验3分别以SCR、VPA及SCR+VPA处理化学激活后的体细胞核移植克隆胚胎,研究并探讨了SCR和VPA联合处理对克隆胚胎的体外发育力的影响。结果表明250nM SCR和2mM VPA联合处理24h组与对照组相比克隆胚胎的囊胚率显著提高(18.3vs9.2%,p<0.05),但与250nM SCR处理24h组(19.9%)及2mM VPA处理24h组(18.9%)比较囊胚率无显著性差异(p>0.05)。实验4研究了SCR对体细胞核移植克隆胚胎的体内发育情况的影响,将250nM SCR处理24h与对照组经过体细胞核移植后获得克隆胚胎分别移植于代孕母后,对照组移植的受体可成功产下健康的克隆仔猪,而SCR处理组并未获得克隆仔猪。综合以上结果表明,SCR处理体细胞核移植胚胎可以提高克隆胚胎的体外囊胚率,对于体内发育能力有必要进一步研究。此外,SCR与VPA联合处理对体细胞核移植克隆胚胎的体外发育无协同作用。
第三章：增强型绿色荧光蛋白(EGFP)在转基因动物的研究中可用来标记目的基因、筛选阳性胚胎、建立荧光动物模型：如将EGFP或红色荧光蛋白基因插入到猪的基因组内,可以很方便的对细胞和组织内某些因子的表达和代谢进行跟踪,从而为基础性研究提供便利。本试验以获得的转EGFP基因阳性细胞作为供体细胞,经核移植和电融合后在2mM VPA处理24h的转基因重构胚移入去核的卵母细胞中,成功地获得了转基因囊胚,并比较克隆胚胎体外及体内发育情况。结果表明GFP转基因重构胚重构胚体外囊胚率为15.1%,与未转基因组11.4%相比无显差异著(P>0.05)。将1～8细胞阶段重构胚胎移植到3头自然发情的五指山小型猪输卵管内,2头未发育到期而返情,1头怀孕106天后,成功产下4头克隆猪。其中2头为死胎,体重分别为247g和313g,其余2头在分娩后由于饲养管理不善死亡,体重分别为366g和573g。经过对猪皮肤组织外来基因检测及受体母猪、克隆猪与供体细胞的13个多态性位点的微卫星检测结果表明,其中2头具有绿色荧光遗传特征,4头克隆猪个体与供核细胞具有完全相同的多态性,而克隆猪个体与代孕母猪的多态性不同,证明克隆仔猪与代孕母猪无亲缘关系。研究结果为转基因五指山小型猪体细胞克隆的技术体系建立提供参考,从而为人类疾病发病机理研究以及生产人类遗传疾病的基因模型动物奠定基础。
Epigenetic modification influences reprogramming and subsequent development of somatic cell nuclear transfer(SCNT)embryos. Such modification includes an increase in histone acetylation. Histone deacetylase inhibitors(HDACi), such as valproic acid (VPA) and scriptaid (SCR), have been known to maintain a high cellular level of histone acetylation. Hence, treatment of nuclear transfer embryos with HDACi may increase the efficiency of cloning. The present study attempted to examine the effects of VPA and SCR with regard to the potency of enhancement of in vitro and in vivo development of Wuzhishan miniature pig SCNT embryos.
In CHAPTER1. The aim of the present study was to examine the effects of VPA, a HDACi on in vitro and in vivo development of Wuzhishan miniature pig SCNT embryos. Experiment1compared the in vitro developmental competence of nuclear transfer embryos treated with various concentrations of VPA for24h. Embryos treated with2mM VPA for24h reported a significantly increased rate of blastocyst formation compared with controls or4mM VPA and8mM VPA treated groups (21.5vs10.5,12.6and17.2%). Experiment2examined the in vitro developmental competence of nuclear transfer embryos treated with2mM VPA for different time periods following chemical activation. Embryos treated for24h reported higher rates of blastocyst formation than the controls or treated for4h and48h groups (20.7vs9.2,12.1and9.1%). In experiment3, an average of207(192to216) nuclear transfer embryos from the VPA-treated group were transferred to surrogate mothers, resulting in three pregnancies. Two of the surrogates delivered a total of11live piglets, the nine of11piglets in the VPA-treated group died in an unknown reason within1day to5days after birch. Untreated control embryos (average,205(179to225)) transferred to four surrogate mothers resulted in three pregnancies, two of which delivered a total of12live offspring, four of the12piglets in the VPA-untreated group died in an unknown reason within1day to3days, eight of12of the12piglets in the VPA-untreated group have survived over3or4months. The average birth weight of the two litters from the VPA-treated group was tended to be lower than that from the control groups (551.6g vs675.2g). These results suggest that VPA treatment increases the rate of blastocyst formation for SCNT embryos, both the VPA-treated and the untreated clones can develop to term in pigs, but the offspring from VPA-treated embryos survived to adulthood lower than the offspring from VPA-treated embryos(18.2vs66.9%; P<0.05).
In CHAPTER2. The aim of the present study was to examine the effects of SCR, a HDACi, on the in vitro and in vivo development of Wuzhishan miniature pig SCNT embryos. Experiment1compared the in vitro developmental competence of nuclear transfer embryos treated with various concentrations of SCR for24h. Embryos treated with250nM SCR showed a significantly increased rate of blastocyst formation compared with controls or embryos treated with50nM or500nM SCR (20.0vs12.2,8.9, and11.9%). Experiment2examined the in vitro developmental competence of nuclear transfer embryos treated with250nM SCR for different time periods following chemical activation. Embryos treated for24h showed higher rates of blastocyst formation than controls or embryos treated for12h or48h (22.2vs11.5,10.0, and18.5%). Experiment3directly compared the effects of SCR and VPA treatment and examined the additive effect of SCR and VPA on nuclear transfer embryos following chemical activation. Embryos treated with250nM SCR or2mM VPA for24h showed a significantly increased rate of blastocyst formation compared with controls (19.9and18.3vs9.2%), but not increased with a combination of250nM SCR and2mM VPA for24h (18.9vs19.9,18.3%). In experiment4, nuclear transfer embryos treated with250nM SCR for24h and non-treated controls were transferred to surrogate mothers, resulting in the birth of healthy cloned pups from the controls but no births in the SCR-treated group. These results suggest that SCR treatment increases the rate of blastocyst formation in somatic cell nuclear transfer embryos and affects their subsequent growth. Also, co-treatment with VPA and SCR did not improve the blastocyst formation rate.
In CHAPTER3. Genetically engineered pigs with cell markers such as fluorescent proteins are highly useful in lines of research that include the tracking of transplanted cells or tissues. It has been demonstrated that increased histone acetylation in SCNT embryos, by applying a histone deacetylase(HDAC) inhibitor such as VPA significantly enhances the developmental competence in our previous research. Herein, we report that the re constructed embryos treatment with2mM VPA for24h, and compared the development of SCNT embryos to the blastocyst stage when Wuzhishan miniature pig ear fibroblasts as donor cells, there have no significant differences were observed in the cleavage rates and blastocyste formation rates in non-trasfected or transfected ear fibroblasts (72.6,15.1and74.5,11.4%). we used the ear fibroblasts of miniature pigs transfected with EGFP as the donor cells, and transplanted reconstructed embryos (at the1to8cell stage) into the oviducts of three sexually mature Wuzhishan miniature pigs that were naturally estrus. One recipient successfully birthed two stillborn and two live piglets after116days of pregnancy. We investigated the presence of the foreign gene in the skin tissues of the piglets and tested13microsatellite polymorphisms in the recipient pigs, cloned offspring and donor cells. The results indicated that two cloned offspring expressed EGFP and all four cloned offspring had the same polymorphisms at all13microsatellites as the donor cell. However, the microsatellites varied between the cloned offspring and the recipient pigs, indicating the absence of a blood relationship between the two. The study therefore establishes a system to obtain transgenic Wuzhishan inbred miniature pigs by SCNT, thereby providing a basis to study the pathologic mechanisms of human disease and producing animal models for human genetic disorders.
引文
[1]Prather RS, Hawley RJ, Carter DB, Lai L, Greenstein JL. Transgenic swine for biomedicine and agriculture. Theriogenology 2003;59:115-123.
[2]Yanbin Z, Chuan L, Chong F, Dengke P, Yuzhu L. Improving the Developmental Potency of Cloned Embryos and Cloning Efficiency in Wuzhishan Inbred Pig by Scriptaid Treatment. Genomics and Applied Biology 2011;30:268-273.
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