小分子化合物对猪胚胎体外发育的影响及iPS嵌合胚胎的研究
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摘要
尽管各种核移植动物已相继诞生,但是其核移植效率并不高。目前,核移植领域的研究主要集中于其重编程分子机理等方面,尤其是研究各种小分子化合物如何提高核移植效率成为该领域研究的热点。同时,重编程技术的另外一种方法,是最近iPS技术的出现,各种类型的iPS细胞已经成功诱导,但是有关检验iPS细胞与胚胎干细胞类似全能性的嵌合体研究,目前却只有在小鼠获得成功。为此,作为重要模式动物和异种器官来源的猪有关iPS细胞嵌合体的研究具有重要的意义。本研究选择研究两种小分子化合物胰岛素-转铁蛋白-硒(ITS)和维生素C(Vc)对猪胚胎体外发育的影响,在优化胚胎体外培养体系基础上,研究影响猪iPS嵌合胚胎的因素,试验结果如下。
1.ITS对猪卵母细胞体外成熟培养和胚胎发育的影响
(1)在猪卵母细胞体外成熟培养中,其添加激素培养42h实验组体外成熟率显著高于对照组(47.8% vs 11.7%, P<0.05),当激素添加培养减少到21h时,第一阶段添加激素(前21h)和第二阶段添加激素(后21h)实验组的体外成熟率显著高于对照组(45.54%和44.8% vs 11.7%, P<0.05);
(2)为了进一步提高卵母细胞成熟率(核成熟),在培养液中添加1%的ITS,实验组卵母细胞体外成熟率显著高于对照组(78.6% vs 54.4%, P<0.05);
(3)对卵母细胞的胞质成熟进一步检测发现,实验组能够显著减少胞质中I和II型皮质颗粒分布(I和II型CGs),同时能够显著提高II型CGs的分布,达85.3%;
(4)对成熟卵母细胞进一步激活,其胚胎体外发育结果表明,实验组和对照组在囊胚发育率方面无显著差异,但实验组的平均囊胚细胞数显著多于对照组(54.56 vs 37.35, P<0.05),说明ITS的添加有利于胚胎的体外发育。由此可见,添加ITS能够使卵母细胞核与质同步成熟率达到67.04% (78.56% x 85.33%),显著高于无ITS添加组34.10%(54.4% x 62.69%),同时能够促进后续胚胎发育的质量。因此,本研究确定的最佳猪卵母细胞体外成熟液为(mM199 + 10 ng/ml EGF + 10IU/ml PMSG + 10IU/ml hCG + 2.5IU/ml FSH+ 1%ITS)。
2.维生素C(Vitamin c,Vc)对猪胚胎体外发育的影响
(1)添加Vc实验组中胚胎内的ROS水平显著低于对照组,在各个添加Vc实验组中以处理E最能显著降低胚胎内ROS水平;
(2)在胚胎发育方面,处理组E的囊胚率(27.74%)显著高于其它处理组(A,10.46%; B, 17.63%; C, 15.64%; D, 17.43%; F, 17.26%; P<0.05),但是,在卵裂率上无显著性差异(P>0.05)。处理组E的平均囊胚细胞数(62.33)显著高于处理组A和B(34.6和46.67, P<0.05),但是,在处理C,D,E和F之间无显著性差异(52.67, 58 , 62.33和56.67) (P<0.05);
(3)对凋亡基因表达的研究,处理组E中囊胚的Bcl-xl与Bax基因表达的相对比值显著高于其它处理(P<0.05),是对照组的4倍;
(4)用TUNEL方法对胚胎内凋亡检测发现,处理组C和E的胚胎发生凋亡细胞数比例(12.21%, 11.94%)显著低于实验组A(23.17%); (5)检测实验组E和F中的多能性基因Nanog的表达,两者之间无显著性差异(P>0.05),但是显著高于其它实验组(P<0.05),其表达量接近于对照组的5倍。由此可见,通过在PZM-3中添加Vc(20μg/ml浓度最佳)能够通过减少胚胎中活性氧的产生,提高猪胚胎在体外发育的效率与质量。
3.不同超排方案对陕北八眉猪超数排卵的影响
(1)用1000IU的eCG进行超排处理与对照组(自然发情)相比,能够提高卵巢的反应效率,产生更多的黄体数(14.3 vs 9.2, P>0.05);
(2)FSH递减剂量的超排方法与eCG超排方法相比,在不同剂量的FSH实验组中都能够显著提高卵巢的反应效率(处理B黄体数: 77.8,处理C黄体数: 66.8和处理D黄体数: 42.7 Vs处理A黄体数: 14.3, P<0.05),但是在处理B和C中有卵巢囊肿的发生,而处理D未发生卵巢囊肿。
上述结果表明,处理组D(FSH:280IU)对于八眉猪是一种合适的超排方法,能够获得较多的体内卵母细胞或胚胎数。由此可见,对八眉猪最合适的该超排方案为:(FSH:280IU; D13/100 IU, D14/80 IU, D15/60 IU, D16/40 IU + PG 0.2 mg, D17/hCG 1000 IU)。4.猪iPS细胞嵌合胚胎创制的研究(1)由iPS-1和iPS-2细胞进行的重构胚胎能够发育到囊胚阶段,但是iPS-3细胞做的重构胚不能有效发育到囊胚阶段,同时也不能有效参与孤雌囊胚及体内囊胚的嵌合;
(2)选择在配种后4.5-5天冲胚,获得囊胚率(37.71%)显著高于在短于4.5天获得的囊胚率(4.56%, P<0.05),同时离体冲胚与手术冲胚方法,在其冲胚率上无显著性差异(83.33% vs 85.92%, P>0.05);
(3)在受体移植方面,其中在移植胚胎数在20枚以上妊娠效果好。为此,在初步进行胚胎学选择iPS细胞后,同时对获得胚胎及受体移植方面进行条件优化,期望通过这几个方面的优化,为后续真正获得猪iPS嵌合体奠定基础。
Although many types of somatic cloning animal successfully have been produced, the efficiency of nuclear transfer is still low. The research in nuclear transfer fields mainly focus on molecular mechanism in nuclear reprogramming, and especially all kinds of small chemical compounds are researched in improving the efficiency. Many kinds of iPS cells are successfully induced after using iPS technique as the other kind of nuclear reprogramming methods, but it is a problem that the chimera to prove the totipotency of iPSc similar to embryonic stem cells is only successfully applied in mouse. Therefore, it is very important to research in producing porcine chimera, which will be a model for human disease and xenotransplantation in future. This study is to research in the effects of two kinds of small chemical compounds (ITS and Vitamin C) on the development of porcine embryos in vitro and analyze the factors of affecting to producing porcine chimera using porcine iPSc, and the experimental restuls shown as following.
1. The effects of insulin–tranferrin–selenium (ITS) on nuclear and cytoplasmic maturation of porcine oocytes and subsequent embryonic development were researched
(1) The rate of oocyte in vitro maturation (IVM) in experimental group treated with hormones for 42 h was significantly increased than that in control group without hormones treatment (47.8% vs. 11.7%, respectively, P<0.05). When reduced the hormone treatment period from 42 h to 21 h, which included both the first 21 h period of hormones treatment (45.4%) and the second 21 h period of hormones treatment (44.8%), the rate of oocyte IVM was still higher than that of control group (P<0.05).
(2) To improve porcine oocyte nuclear maturation, 1% ITS was added in medium supplemented with hormones. The rate of nuclear maturation in ITS treated group was significantly higher than ITS untreated group (78.6% vs 54.4%, respectively, P<0.05).
(3) ITS treatment was also significantly reduced the ratio of oocytes with type I and type III CGs distributions, and significantly increased the ratio of oocytes with type II CGs distribution (85.3%). The results indicated that synchronization rates of nuclear and ooplasmic maturation reached to 67.04% (78.56% x 85.33%).
In conclusion, the group of modified Tissue Culture Medium 199 (mM199) + 10 ng/ml EGF + 10 IU/ml pregnant mare serum gonadotrophin (PMSG) + 10 IU/ml human chorion gonadotrophin (hCG) + 2.5 IU/ml follicle stimulate hormone (FSH) + 1%ITS is suitable for culturing porcine oocytes in vitro.
2. The effects of Vc on development of porcine embryos were researched in vitro
In order to clear out the effects of Vc (0, 2.5, 5, 10, 20, 40μg/ml; group A, B, C, D, E, F) adding in PZM-3 were choosed to culture parthenogenetic presumptive diploid zygotes produced by electroactivation after oocytes maturation in vitro.
(1)the ROS level in embryos significantly decreased in the Vc adding groups compared to the group A (the control group ),and the group E is significantly decreased ROS level in embryos compared to other groups with adding Vc(Group B, C, D, F).
(2)The Rate of blastocyst in group E (27.74%) is significantly higher than in other groups (P<0.05) but there was not significant differences in the rate of embryo cleavage in different groups(P>0.05), and the total nuclei number in blastocyst of Group E(62.33) is significantly higher than Group A and B(34.67 Vs 46.67, P<0.05), but there was not significant difference in Group C,D,E and F(52.67, 58, 62.33 and 56.67; P>0.05).
(3)The relative expression of Bcl-xl to Bax gene in Group E is significantly higher than other groups (P<0.05), and it was almost 4 folds to the control.
(4) It is showed that TUNEL index in adding Vc groups (Group C and E: 12.21% and 11.94%) significantly decreased compared to nonsupplemented Group (Conrol: 23.17%, P<0.05) through analyzing TUNEL results (Group A, C, E).
(5) The expression of Nanog gene in Group E and F is significantly higher than other groups (P<0.05) but there is not significant difference in this two groups (P>0.05) and Group E and F was almost 5 folds to the control.
These data thus suggest that Vc , which 20μg/ml in PZM-3 was the best suitable dosage, improves the development rate and quality of porcine parthenotes by preventing oxidative damage in the process of porcine embryo development in vitro.
3. The effects of different superovulation procotols on ovary reaction of Baimei gilts were researched
The objective of this study was to compare the superovulation effects between equine chorionic gonadotropin (eCG , Group A) and FSH (Group B-D) in Chinese Bamei gilts.
(1) The results show that though eCG could produce more corpus lutea (CL, 14.3) than the control (CL, 9.2), the FSH treatments were significantly increased the number of CL than that of eCG treatment.
(2) Within the different FSH protocols, the numbers of CL were significantly more in Group B (CL, 77.8) and C (CL, 66.8) than that in Group D (CL, 42.7), however, ovarian cysts were observed in Group B and C, but not in Group D. These data suggest that Group D (280IU FSH) is a suitable protocol to facilitate the development of ovarian follicles and may obtain more useful embryos per gilt through embryos recovery.
The optimal FSH protocol of superovulation in Bamei gilts is: D13/100 IU, D14/80 IU, D15/60 IU, D16/40 IU plus PG 0.2 mg, D17/hCG 1000 IU
4. Research on creating chimera embryos using porcine iPS cells as donor cells
The research of producing gilts of iPSc chimera is very important because it has been a model for human disease and xenotransplantation in future. The objective of this study is to choose suitable iPSc for chimera embryo transfer through analyzing the development of reconstructed embryos using iPSc as donor cells through nuclear transfer and chimera embros in vitro before transferring embryos into gilts, and research in time and methods of obtaining the porcine embryos in vivo.
(1)The nuclear transfer embryos cannot develop to blastocyte stage using iPS-3 as donor cells and the iPS-3 cell cannot involve in the development of chimera embryos through using blasocytes from in vitro and vivo, the ratio of blastocyte in reconstructed embryos development usig iPS-1 and iPS-3(6.98%, 8.07%) as donor cells is significantly lower than the control(13.16%,P<0.05).
(2) The ratio of blastocystes in recovering embryos between 4.5 to 5 days(37.71%) is significantly higher than that below 4.5days after first mating(4.56%, P<0.05),and there is not significant difference in the rate of recovering porcine embryo between surgical and in vitro uterus methods(83.33% vs 85.92%, P>0.05).
Therefore, the system of choosing suitable iPSc for donor cells and obtaining many porcine embryos with good quality is basically built for the next porcine iPSc chimera research.
1.ITS对猪卵母细胞体外成熟培养和胚胎发育的影响
(1)在猪卵母细胞体外成熟培养中,其添加激素培养42h实验组体外成熟率显著高于对照组(47.8% vs 11.7%, P<0.05),当激素添加培养减少到21h时,第一阶段添加激素(前21h)和第二阶段添加激素(后21h)实验组的体外成熟率显著高于对照组(45.54%和44.8% vs 11.7%, P<0.05);
(2)为了进一步提高卵母细胞成熟率(核成熟),在培养液中添加1%的ITS,实验组卵母细胞体外成熟率显著高于对照组(78.6% vs 54.4%, P<0.05);
(3)对卵母细胞的胞质成熟进一步检测发现,实验组能够显著减少胞质中I和II型皮质颗粒分布(I和II型CGs),同时能够显著提高II型CGs的分布,达85.3%;
(4)对成熟卵母细胞进一步激活,其胚胎体外发育结果表明,实验组和对照组在囊胚发育率方面无显著差异,但实验组的平均囊胚细胞数显著多于对照组(54.56 vs 37.35, P<0.05),说明ITS的添加有利于胚胎的体外发育。由此可见,添加ITS能够使卵母细胞核与质同步成熟率达到67.04% (78.56% x 85.33%),显著高于无ITS添加组34.10%(54.4% x 62.69%),同时能够促进后续胚胎发育的质量。因此,本研究确定的最佳猪卵母细胞体外成熟液为(mM199 + 10 ng/ml EGF + 10IU/ml PMSG + 10IU/ml hCG + 2.5IU/ml FSH+ 1%ITS)。
2.维生素C(Vitamin c,Vc)对猪胚胎体外发育的影响
(1)添加Vc实验组中胚胎内的ROS水平显著低于对照组,在各个添加Vc实验组中以处理E最能显著降低胚胎内ROS水平;
(2)在胚胎发育方面,处理组E的囊胚率(27.74%)显著高于其它处理组(A,10.46%; B, 17.63%; C, 15.64%; D, 17.43%; F, 17.26%; P<0.05),但是,在卵裂率上无显著性差异(P>0.05)。处理组E的平均囊胚细胞数(62.33)显著高于处理组A和B(34.6和46.67, P<0.05),但是,在处理C,D,E和F之间无显著性差异(52.67, 58 , 62.33和56.67) (P<0.05);
(3)对凋亡基因表达的研究,处理组E中囊胚的Bcl-xl与Bax基因表达的相对比值显著高于其它处理(P<0.05),是对照组的4倍;
(4)用TUNEL方法对胚胎内凋亡检测发现,处理组C和E的胚胎发生凋亡细胞数比例(12.21%, 11.94%)显著低于实验组A(23.17%); (5)检测实验组E和F中的多能性基因Nanog的表达,两者之间无显著性差异(P>0.05),但是显著高于其它实验组(P<0.05),其表达量接近于对照组的5倍。由此可见,通过在PZM-3中添加Vc(20μg/ml浓度最佳)能够通过减少胚胎中活性氧的产生,提高猪胚胎在体外发育的效率与质量。
3.不同超排方案对陕北八眉猪超数排卵的影响
(1)用1000IU的eCG进行超排处理与对照组(自然发情)相比,能够提高卵巢的反应效率,产生更多的黄体数(14.3 vs 9.2, P>0.05);
(2)FSH递减剂量的超排方法与eCG超排方法相比,在不同剂量的FSH实验组中都能够显著提高卵巢的反应效率(处理B黄体数: 77.8,处理C黄体数: 66.8和处理D黄体数: 42.7 Vs处理A黄体数: 14.3, P<0.05),但是在处理B和C中有卵巢囊肿的发生,而处理D未发生卵巢囊肿。
上述结果表明,处理组D(FSH:280IU)对于八眉猪是一种合适的超排方法,能够获得较多的体内卵母细胞或胚胎数。由此可见,对八眉猪最合适的该超排方案为:(FSH:280IU; D13/100 IU, D14/80 IU, D15/60 IU, D16/40 IU + PG 0.2 mg, D17/hCG 1000 IU)。4.猪iPS细胞嵌合胚胎创制的研究(1)由iPS-1和iPS-2细胞进行的重构胚胎能够发育到囊胚阶段,但是iPS-3细胞做的重构胚不能有效发育到囊胚阶段,同时也不能有效参与孤雌囊胚及体内囊胚的嵌合;
(2)选择在配种后4.5-5天冲胚,获得囊胚率(37.71%)显著高于在短于4.5天获得的囊胚率(4.56%, P<0.05),同时离体冲胚与手术冲胚方法,在其冲胚率上无显著性差异(83.33% vs 85.92%, P>0.05);
(3)在受体移植方面,其中在移植胚胎数在20枚以上妊娠效果好。为此,在初步进行胚胎学选择iPS细胞后,同时对获得胚胎及受体移植方面进行条件优化,期望通过这几个方面的优化,为后续真正获得猪iPS嵌合体奠定基础。
Although many types of somatic cloning animal successfully have been produced, the efficiency of nuclear transfer is still low. The research in nuclear transfer fields mainly focus on molecular mechanism in nuclear reprogramming, and especially all kinds of small chemical compounds are researched in improving the efficiency. Many kinds of iPS cells are successfully induced after using iPS technique as the other kind of nuclear reprogramming methods, but it is a problem that the chimera to prove the totipotency of iPSc similar to embryonic stem cells is only successfully applied in mouse. Therefore, it is very important to research in producing porcine chimera, which will be a model for human disease and xenotransplantation in future. This study is to research in the effects of two kinds of small chemical compounds (ITS and Vitamin C) on the development of porcine embryos in vitro and analyze the factors of affecting to producing porcine chimera using porcine iPSc, and the experimental restuls shown as following.
1. The effects of insulin–tranferrin–selenium (ITS) on nuclear and cytoplasmic maturation of porcine oocytes and subsequent embryonic development were researched
(1) The rate of oocyte in vitro maturation (IVM) in experimental group treated with hormones for 42 h was significantly increased than that in control group without hormones treatment (47.8% vs. 11.7%, respectively, P<0.05). When reduced the hormone treatment period from 42 h to 21 h, which included both the first 21 h period of hormones treatment (45.4%) and the second 21 h period of hormones treatment (44.8%), the rate of oocyte IVM was still higher than that of control group (P<0.05).
(2) To improve porcine oocyte nuclear maturation, 1% ITS was added in medium supplemented with hormones. The rate of nuclear maturation in ITS treated group was significantly higher than ITS untreated group (78.6% vs 54.4%, respectively, P<0.05).
(3) ITS treatment was also significantly reduced the ratio of oocytes with type I and type III CGs distributions, and significantly increased the ratio of oocytes with type II CGs distribution (85.3%). The results indicated that synchronization rates of nuclear and ooplasmic maturation reached to 67.04% (78.56% x 85.33%).
In conclusion, the group of modified Tissue Culture Medium 199 (mM199) + 10 ng/ml EGF + 10 IU/ml pregnant mare serum gonadotrophin (PMSG) + 10 IU/ml human chorion gonadotrophin (hCG) + 2.5 IU/ml follicle stimulate hormone (FSH) + 1%ITS is suitable for culturing porcine oocytes in vitro.
2. The effects of Vc on development of porcine embryos were researched in vitro
In order to clear out the effects of Vc (0, 2.5, 5, 10, 20, 40μg/ml; group A, B, C, D, E, F) adding in PZM-3 were choosed to culture parthenogenetic presumptive diploid zygotes produced by electroactivation after oocytes maturation in vitro.
(1)the ROS level in embryos significantly decreased in the Vc adding groups compared to the group A (the control group ),and the group E is significantly decreased ROS level in embryos compared to other groups with adding Vc(Group B, C, D, F).
(2)The Rate of blastocyst in group E (27.74%) is significantly higher than in other groups (P<0.05) but there was not significant differences in the rate of embryo cleavage in different groups(P>0.05), and the total nuclei number in blastocyst of Group E(62.33) is significantly higher than Group A and B(34.67 Vs 46.67, P<0.05), but there was not significant difference in Group C,D,E and F(52.67, 58, 62.33 and 56.67; P>0.05).
(3)The relative expression of Bcl-xl to Bax gene in Group E is significantly higher than other groups (P<0.05), and it was almost 4 folds to the control.
(4) It is showed that TUNEL index in adding Vc groups (Group C and E: 12.21% and 11.94%) significantly decreased compared to nonsupplemented Group (Conrol: 23.17%, P<0.05) through analyzing TUNEL results (Group A, C, E).
(5) The expression of Nanog gene in Group E and F is significantly higher than other groups (P<0.05) but there is not significant difference in this two groups (P>0.05) and Group E and F was almost 5 folds to the control.
These data thus suggest that Vc , which 20μg/ml in PZM-3 was the best suitable dosage, improves the development rate and quality of porcine parthenotes by preventing oxidative damage in the process of porcine embryo development in vitro.
3. The effects of different superovulation procotols on ovary reaction of Baimei gilts were researched
The objective of this study was to compare the superovulation effects between equine chorionic gonadotropin (eCG , Group A) and FSH (Group B-D) in Chinese Bamei gilts.
(1) The results show that though eCG could produce more corpus lutea (CL, 14.3) than the control (CL, 9.2), the FSH treatments were significantly increased the number of CL than that of eCG treatment.
(2) Within the different FSH protocols, the numbers of CL were significantly more in Group B (CL, 77.8) and C (CL, 66.8) than that in Group D (CL, 42.7), however, ovarian cysts were observed in Group B and C, but not in Group D. These data suggest that Group D (280IU FSH) is a suitable protocol to facilitate the development of ovarian follicles and may obtain more useful embryos per gilt through embryos recovery.
The optimal FSH protocol of superovulation in Bamei gilts is: D13/100 IU, D14/80 IU, D15/60 IU, D16/40 IU plus PG 0.2 mg, D17/hCG 1000 IU
4. Research on creating chimera embryos using porcine iPS cells as donor cells
The research of producing gilts of iPSc chimera is very important because it has been a model for human disease and xenotransplantation in future. The objective of this study is to choose suitable iPSc for chimera embryo transfer through analyzing the development of reconstructed embryos using iPSc as donor cells through nuclear transfer and chimera embros in vitro before transferring embryos into gilts, and research in time and methods of obtaining the porcine embryos in vivo.
(1)The nuclear transfer embryos cannot develop to blastocyte stage using iPS-3 as donor cells and the iPS-3 cell cannot involve in the development of chimera embryos through using blasocytes from in vitro and vivo, the ratio of blastocyte in reconstructed embryos development usig iPS-1 and iPS-3(6.98%, 8.07%) as donor cells is significantly lower than the control(13.16%,P<0.05).
(2) The ratio of blastocystes in recovering embryos between 4.5 to 5 days(37.71%) is significantly higher than that below 4.5days after first mating(4.56%, P<0.05),and there is not significant difference in the rate of recovering porcine embryo between surgical and in vitro uterus methods(83.33% vs 85.92%, P>0.05).
Therefore, the system of choosing suitable iPSc for donor cells and obtaining many porcine embryos with good quality is basically built for the next porcine iPSc chimera research.
引文
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