维甲酸对鸡胚生殖细胞增殖和减数分裂调控的研究
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摘要
家禽的胚胎发育有独特的规律,通过对家禽原始生殖细胞(primordial germ cell, PGC)的体外研究,可以更深入地了解其胚胎发育过程及影响因素,并可对其基因进行遗传操作,制作各种研究模型,从而为生殖生物学和发育生物学等研究提供了一个优良的动物模型。为了更深层次揭示禽类生殖细胞增殖和分化的细胞内部和外源性调控机理、性腺发育和配子形成的分子机理,本实验以海兰鸡鸡胚为材料,分离了不同时期的PGC,进行体外培养并优化培养模型,研究了维甲酸(RA)对PGC粘附和增殖的作用及其胞内信号传导机制,同时还研究了RA对胚胎期卵巢生殖细胞减数分裂启动过程中的调控作用,探索基于LV-siRNA的慢病毒载体法生产转基因家禽的可行性,建立Raldh2基因敲除的转基因鸡模型,为进一步研究胚胎早期发育的机理、转基因家禽的制备奠定了基础。
1.鸡胚PGC的分离、培养及鉴定
在体视显微镜下用玻璃针分离19期3.5d鸡胚生殖嵴部位或28期5.5d鸡胚性腺,胰蛋白酶+EDTA消化分散组织,用KO-DMEM添加7.5%胎牛血清和10ng/ml白血病抑制因子(LIF)、10ng/ml碱性成纤维生长因子(bFGF)、10ng/ml干细胞生长因子(SCF)、0.1mMMEM非必需氨基酸、0.1mMβ-巯基乙醇、2mM L-谷氨酰胺(Glu)、100IU/ml青霉素和100μg/ml链霉素的培养液,通过PGC和体细胞体外共培养的方式建立原代培养模型。然后传代于鸡胚成纤维细胞饲养层上进行继代培养。同时我们从13-15期鸡胚血液中分离提取PGC,在饲养层上进行原代和传代培养。传至3-5代后,用碱性磷酸酶(AKP)、过碘酸雪夫氏(PAS)化学反应及c-kit、SSEA-1,3,4、EMA-1等免疫细胞化学鉴定PGC,用PCNA和BrdU掺入法检测细胞的增殖活性。并收集PGC集落,采用RT-PCR方法检测干细胞多能性相关基因(Pou5fl,Sox2和Nanog)的表达。上述结果表明我们建立的PGC-体细胞共培养模型较好的维持了PGC的生物特性及未分化状态,且具有较强的增殖活性。因此,该模型是可用于PGC体外增殖和分化调控的研究。
2.RA对鸡胚PGC粘附和增殖的影响及其机理的研究
本实验研究了RA对鸡胚PGC粘附和增殖的影响及其相关的信号传导途径。结果表明：经过RA(0.1-10μM)处理后,PGC的集落数目和面积呈时间和剂量依赖性增加。同时RA能够促进细胞间的粘附,提高粘附蛋白E-钙粘蛋白、α-连锁蛋白和β-连锁蛋白的mRNA水平,增强PGC的粘附性,并存在一定的剂量效应关系。流式分析结果显示,PGC经过RA处理4天后,S/G2期(4N)的PGC数量显著增加,说明此部分细胞已经进入有丝分裂期。此外,RA能激活PGC中PKC和Akt活性,促进NF-κB的核转移,提高钙粘蛋白和β-catenin的磷酸化水平,然而这些刺激作用分别被LY294002(P13K抑制剂),KP372-1(Akt抑制剂),SN50(NF-κB抑制剂)和H7(PKC抑制剂)所阻断。RA显著上调PGC中细胞周期蛋白cyclin D1/E, CDK2/6和原癌基因c-fos, c-myc(?)勺表达,但是这种上调作用被H7、LY294002、KP372-1和SN50所抑制：综上所述,在体外培养条件下,RA通过PI3K/Akt/NF-κB信号级联反应及与PKC/β-catenin之间的交联反应促进鸡胚PGC的增殖,揭示RA在调控鸡胚胎生殖细胞发育中起着重要的作用。
3.RA对鸡胚生殖细胞减数分裂的调控机理的研究
在本实验中,我们结合胚胎期卵巢的体外培养和RNA干扰等手段,探索RA对鸡胚胎期生殖细胞减数分裂的调控及作用机制。SCP3和γH2AX免疫荧光染色结果表明,鸡胚卵巢生殖细胞在15.5天进入减数分裂,比例逐渐增加,到18.5天,进入减数分裂的细胞几乎遍布整个卵巢。减数分裂早期标志基因Strs8mRNA的表达在12.5天开始显著上调,早于减数分裂启动的时间：而减数分裂标志基因Scp3和Dmcl mRNA(?)内表达则在15.5天显著上调,再次验证鸡胚卵巢生殖细胞在15.5天进入减数分裂；RA合成酶基因Raldh2在15.5天显著上调而降解酶Cyp26bl却逐渐下降,表明Raldh2在RA聚集并维持减数分裂中的重要作用；RA受体RARβ则略微升高,表明其在RA启动减数分裂中的介导作用。而在雄性睾丸中,这些基因的表达变化很小,一直保持较低水平。10.5天和12.5天鸡胚卵巢在无血清的体外培养体系中分别培养5天和3天,生殖细胞能白发启动减数分裂：在培养体系中添加RA能明显增加进入减数分裂的细胞比例,显著上调了减数分裂相关基因Stra8,Scp3和Dmcl(?)的mRNA表达水平。Raldh2shRNA干扰后显著抑制了RA诱导减数分裂启动的作用。综上所述,我们的研究表明,RA及其信号通路对减数分裂启动中的重要调控作用,并首次报道了Raldh2基因敲减能够显著抑制生殖细胞进入减数分裂。
4.利用shRNA慢病毒载体制备转基因鸡模型研究生殖细胞的发育
本实验采用RNA干涉(RNAi)的方法,成功设计了三对针对Raldh2基因序列的保守区域的shRNA (short hairpin)分子,并构建质粒干涉载体。转染鸡胚15.5天卵巢生殖细胞后,转染效率达到80%-90%。通过qRT-PCR及SCP3免疫荧光染色检测,结果表明Raldh2-gallus-1224很好的抑制了Raldh2的表达,成功筛选出能高效抑制Raldh2复制的靶位点。针对有效位点构建了携带有能高效抑制Raldh2的shRNA慢病毒载体,将包装后的病毒注射到X期鸡胚胚盘内进行转基因操作,建立Raldh2基因敲减的转基因鸡模型。结果发现Raldh2基因敲减的转基因鸡胚外观与对照组的鸡胚相比无明显差异,卵巢形态大致正常,应用PCR检测GFP在胚体内的表达,进一步证实我们成功制备了转基因鸡模型,且qRT-PCR检测到Raldh2的表达显著下降。本实验为探索基于LV-siRNA慢病毒载体法生产转基因家禽的可行性,建立基因敲减的转基因鸡模型,为研究卵泡发育的分子机制奠定基础。
以上实验结果表明：鸡胚PGC的体外共培养模型可用于PGC增殖和分化调控的研究,经AKP、PAS及c-kit、SSEA-1、3、4和EMA-1免疫细胞化学染色和干细胞多能性相关基因(Pou5fl,Sox2和Nanog)表达的RT-PCR检测等多种方法证实了PGC的原始性。利用此模型我们研究了维甲酸在PGC体外培养中的促增殖和粘附作用及其分子机理：我们还描述了鸡胚卵巢生殖细胞体内减数分裂启动中的形态学和分子变化,通过组织培养和RNA干扰等手段研究了RA对减数分裂启动的调控作用。这些发现将为禽类干细胞系的建立和转基因模型的制备奠定基础,同时也为进一步研究生殖细胞的发育提供理论指导和实验平台。
For the development pattern of chicken embryos is unique, we could further understand of the development process and influential factors by study of chicken primordial germ cell (PGC) in vitro. We can acquire many study models by genetic manipulation in PGC of chicken, which can be used as an ideal model for reproductive biology and developmental biology to reveal intrinsic and external molecular mechanisms underlying regulation in proliferation and differentiation of germ cells, gonad development and gametogenesis. In this study, PGC were isolated from Hailan chicken embryo, and then subcultured on feed layer with somatic cells after initial primary culture. In addition, the effect of RA on proliferation of cultured PGC and meiosis initiation of germ cells in embryonic chicken ovary were evaluated, together with underlining mechanisms. Meanwhile, we also investigate the production of transgenic chicken by LV-siRNA and get a model of Raldh2knock down. These studies will provide theoretic guidance for further investigation of early development and preparation of transgenic poultry.
1. Isolation, culture and identification of chicken PGC
PGC were collected from stage14chicken embryonic bloods, stage19genital ridges or stage28gonads with a fine glass needle under a microsurgery scope. For primary culture, cell suspension containing both PGC and somatic cells was seeded onto gelatin-treated6-well culture plates at a density of1×106/well in KO-DMEM supplemented with7.5%fetal calf serum (FCS),10ng/ml leukemia inhibitory factor (LIF),10ng/ml human basic fibroblast growth factor (bFGF),0.1mM MEM nonessential amino acids.0.1mM2-mercaptoethanol,2mM L-glutamine (Glu),100U/ml penicillin and100μg/ml streptomycin, PGC were cultured at38.5℃in an air atmosphere containing5%CO2with60%-70%relative humidity. To trace the origin of the colonies, the primary formed colonies were picked up with a fine glass needle and then subjected to RT-PCR analysis for expression of PGC-specific markers. For further subculture, colonies that were positive for PGC markers were picked up and digested to single cell suspension and reseeded onto6-well dish. The third passage PGC were identified by staining of Alkaline phosphatase (AKP), periodic acid-Schiff regent (PAS), c-kit, stage-specific embryonic antigens (SSEA-1,3.4) and EMA-1immunocytochemistry, and the expression of the pluripotency-associated genes PouV, Nanog and Sox2was analyzed by RT-PCR. The results all confirmed the characteristics of cultured PGC which indicated that the primary and subculture models of PGC could be used for studies about regulation of PGC proliferation and differenciation.
2. Effects of RA on adhesion and proliferation of chicken PGC
In the present study, the proliferating effect of RA on PGC was investigated along with the intracellular PI3K/Akt-mediated NF-κB signaling cascade and PKC/(3-catenin signaling. Results showed that RA significantly promoted PGC proliferation in a dose-and time-dependent manner, which was confirmed by BrdU incorporation. RA induced PGC aggregation by increasing expression of E-cadherin and α/β-catenins. However, this promoting effect was attenuated obviously by sequential inhibitors of LY294002for PI3K, KP372-1, for Akt SN50for NF-κB and PKC inhibitor H7, respectively. Furthermore, Western blot analysis manifested increased Akt phosphorylation (Ser ") of PGC after stimulation with RA, but this effect was abolished by LY294002or KP372-1. Meanwhile, treatment with RA increased expression of NF-κB and decreased IκBα expression that were inhibited by SN50. Moreover, blockade of PI3K or Akt activity resulted in inhibiting NF-κB translocation from the cytoplasm to the nucleus. E-cadherin and β-catenin protein expression levels also were increased by RA addition. Finally, the up-regulated mRNA expression of cell cycle regulating genes (cyclin D1and E, cyclin-dependent kinases6and2) was observed in the RA-treated cells. Flow cytometry analysis confirmed that RA-treated cultured PGC populations displayed a significant increase in the proportion of S and G2phase cells. Again, this stimulation was remarkably retarded by combined treatment with LY294002, KP372-1, SN50and H7respectively. Therefore, these results suggest that RA promote proliferation of the cultured chicken PGC via the PI3K/Akt mediated NF-κB signaling cascade and PKC/β-catenin signaling pathway.
3. Effect of RA on meiosis initiation in the chicken embryo
In the present study, the effect of RA on meiosis initiation in embryonic germ cells was investigated by organ culture in vitro and RNA interference. The results showed the profile of meiosis in chicken embryo ovary-meiotic germ cells were increased gradually since day15.5(first detectable) to day18.5(widespread) by immunofluorescence of SCP3and yH2AX. And the expression of Stra8is specifically up-regulated at day12.5, Scp3and Dmcl became elevated at day15.5. In addition, we observed increased mRNA levels for Raldh2but deceased gradually for Cyp26bl during meiosis that evidence for a requirement of RA accumulation to sustain meiosis. The expression of RARβ also presented slightly increase after day15.5. But in males, the expression of these genes keeps low level throughout development. We also found that the culture ovaries from day10.5and12.5were able to initiate meiosis, and meiosis is stimulated by RA compared with control (P<0.05). Finally, Raldh2shRNA remarkably abolished RA-dependent functions, indicating that RA synthesis and signaling is crucial for meiosis initiation. Taken together, these studies indicate that RA and signal are important in regulating the onset of meiosis in the chicken embryonic ovary. We reported here for the first time that Raldh2knock-down could significantly reduced germ cells to enter meiosis.
4. Production of transgenic chicken via LV-siRNA system
We designed three pairs of shRNA specific for Raldh2by RNA interference to knock down its function and constructed expressing vector of shRNA The day15.5ovarian cells were transfected with either shRNA specific for Raldh2. or a non-targeting shRNA as a negative control. The efficiency of transfection is about80%-90%. Among of these shRNAs, shRNA2produced the most significant inhibitory effect on Raldh2mRNA expression. RNA interference of Raldh2prevented the appearance of meiotic cells by immunofluorescence of SCP3. The Lentivirus vector specific for Raldh2shRNA was constructed and packaged. Then the virus was injected into X stage chicken embryos for production of transgenic chicken. The chicken model of Raldh2knock down was successfully obtained. The results showed that there were no obviously difference between transgenic chicken and wild chicken, as the morphology of ovary. The GFP was detected by PCR, which confirmed the extra gene was injected to genome successfully. The expression of Raldh2was decreased significantly by qRT-PCR. This research provide theoretical basis for understanding the molecular mechanisms of regulation of oogenesis and follicular development.
The above results indicated that the co-culture model of somatic cells and PGC from chicken embryo could be used to study on proliferation and differentiation. PGC were characterized by staining of AKP, PAS, c-kit, SSEA-1,3,4and EMA-1immunocytochemistry. The pluripotency of PGC was also detected by the expression of the pluripotency-associated genes (PouV,Nanog and Sox2). We investigated the molecular mechanisms of RA induced PGC proliferation by this model. Furthermore, we described the morphology and genetic changes during meiosis initiation in chicken embryonic ovary, then studied the regulating effect of RA on meiosis initiation by organ culture in vitro and RNA interference. These findings provide theoretic guidance and experimental platform for studying the development of germ cells and preparation of chimeras and transgenic poultry.
1.鸡胚PGC的分离、培养及鉴定
在体视显微镜下用玻璃针分离19期3.5d鸡胚生殖嵴部位或28期5.5d鸡胚性腺,胰蛋白酶+EDTA消化分散组织,用KO-DMEM添加7.5%胎牛血清和10ng/ml白血病抑制因子(LIF)、10ng/ml碱性成纤维生长因子(bFGF)、10ng/ml干细胞生长因子(SCF)、0.1mMMEM非必需氨基酸、0.1mMβ-巯基乙醇、2mM L-谷氨酰胺(Glu)、100IU/ml青霉素和100μg/ml链霉素的培养液,通过PGC和体细胞体外共培养的方式建立原代培养模型。然后传代于鸡胚成纤维细胞饲养层上进行继代培养。同时我们从13-15期鸡胚血液中分离提取PGC,在饲养层上进行原代和传代培养。传至3-5代后,用碱性磷酸酶(AKP)、过碘酸雪夫氏(PAS)化学反应及c-kit、SSEA-1,3,4、EMA-1等免疫细胞化学鉴定PGC,用PCNA和BrdU掺入法检测细胞的增殖活性。并收集PGC集落,采用RT-PCR方法检测干细胞多能性相关基因(Pou5fl,Sox2和Nanog)的表达。上述结果表明我们建立的PGC-体细胞共培养模型较好的维持了PGC的生物特性及未分化状态,且具有较强的增殖活性。因此,该模型是可用于PGC体外增殖和分化调控的研究。
2.RA对鸡胚PGC粘附和增殖的影响及其机理的研究
本实验研究了RA对鸡胚PGC粘附和增殖的影响及其相关的信号传导途径。结果表明：经过RA(0.1-10μM)处理后,PGC的集落数目和面积呈时间和剂量依赖性增加。同时RA能够促进细胞间的粘附,提高粘附蛋白E-钙粘蛋白、α-连锁蛋白和β-连锁蛋白的mRNA水平,增强PGC的粘附性,并存在一定的剂量效应关系。流式分析结果显示,PGC经过RA处理4天后,S/G2期(4N)的PGC数量显著增加,说明此部分细胞已经进入有丝分裂期。此外,RA能激活PGC中PKC和Akt活性,促进NF-κB的核转移,提高钙粘蛋白和β-catenin的磷酸化水平,然而这些刺激作用分别被LY294002(P13K抑制剂),KP372-1(Akt抑制剂),SN50(NF-κB抑制剂)和H7(PKC抑制剂)所阻断。RA显著上调PGC中细胞周期蛋白cyclin D1/E, CDK2/6和原癌基因c-fos, c-myc(?)勺表达,但是这种上调作用被H7、LY294002、KP372-1和SN50所抑制：综上所述,在体外培养条件下,RA通过PI3K/Akt/NF-κB信号级联反应及与PKC/β-catenin之间的交联反应促进鸡胚PGC的增殖,揭示RA在调控鸡胚胎生殖细胞发育中起着重要的作用。
3.RA对鸡胚生殖细胞减数分裂的调控机理的研究
在本实验中,我们结合胚胎期卵巢的体外培养和RNA干扰等手段,探索RA对鸡胚胎期生殖细胞减数分裂的调控及作用机制。SCP3和γH2AX免疫荧光染色结果表明,鸡胚卵巢生殖细胞在15.5天进入减数分裂,比例逐渐增加,到18.5天,进入减数分裂的细胞几乎遍布整个卵巢。减数分裂早期标志基因Strs8mRNA的表达在12.5天开始显著上调,早于减数分裂启动的时间：而减数分裂标志基因Scp3和Dmcl mRNA(?)内表达则在15.5天显著上调,再次验证鸡胚卵巢生殖细胞在15.5天进入减数分裂；RA合成酶基因Raldh2在15.5天显著上调而降解酶Cyp26bl却逐渐下降,表明Raldh2在RA聚集并维持减数分裂中的重要作用；RA受体RARβ则略微升高,表明其在RA启动减数分裂中的介导作用。而在雄性睾丸中,这些基因的表达变化很小,一直保持较低水平。10.5天和12.5天鸡胚卵巢在无血清的体外培养体系中分别培养5天和3天,生殖细胞能白发启动减数分裂：在培养体系中添加RA能明显增加进入减数分裂的细胞比例,显著上调了减数分裂相关基因Stra8,Scp3和Dmcl(?)的mRNA表达水平。Raldh2shRNA干扰后显著抑制了RA诱导减数分裂启动的作用。综上所述,我们的研究表明,RA及其信号通路对减数分裂启动中的重要调控作用,并首次报道了Raldh2基因敲减能够显著抑制生殖细胞进入减数分裂。
4.利用shRNA慢病毒载体制备转基因鸡模型研究生殖细胞的发育
本实验采用RNA干涉(RNAi)的方法,成功设计了三对针对Raldh2基因序列的保守区域的shRNA (short hairpin)分子,并构建质粒干涉载体。转染鸡胚15.5天卵巢生殖细胞后,转染效率达到80%-90%。通过qRT-PCR及SCP3免疫荧光染色检测,结果表明Raldh2-gallus-1224很好的抑制了Raldh2的表达,成功筛选出能高效抑制Raldh2复制的靶位点。针对有效位点构建了携带有能高效抑制Raldh2的shRNA慢病毒载体,将包装后的病毒注射到X期鸡胚胚盘内进行转基因操作,建立Raldh2基因敲减的转基因鸡模型。结果发现Raldh2基因敲减的转基因鸡胚外观与对照组的鸡胚相比无明显差异,卵巢形态大致正常,应用PCR检测GFP在胚体内的表达,进一步证实我们成功制备了转基因鸡模型,且qRT-PCR检测到Raldh2的表达显著下降。本实验为探索基于LV-siRNA慢病毒载体法生产转基因家禽的可行性,建立基因敲减的转基因鸡模型,为研究卵泡发育的分子机制奠定基础。
以上实验结果表明：鸡胚PGC的体外共培养模型可用于PGC增殖和分化调控的研究,经AKP、PAS及c-kit、SSEA-1、3、4和EMA-1免疫细胞化学染色和干细胞多能性相关基因(Pou5fl,Sox2和Nanog)表达的RT-PCR检测等多种方法证实了PGC的原始性。利用此模型我们研究了维甲酸在PGC体外培养中的促增殖和粘附作用及其分子机理：我们还描述了鸡胚卵巢生殖细胞体内减数分裂启动中的形态学和分子变化,通过组织培养和RNA干扰等手段研究了RA对减数分裂启动的调控作用。这些发现将为禽类干细胞系的建立和转基因模型的制备奠定基础,同时也为进一步研究生殖细胞的发育提供理论指导和实验平台。
For the development pattern of chicken embryos is unique, we could further understand of the development process and influential factors by study of chicken primordial germ cell (PGC) in vitro. We can acquire many study models by genetic manipulation in PGC of chicken, which can be used as an ideal model for reproductive biology and developmental biology to reveal intrinsic and external molecular mechanisms underlying regulation in proliferation and differentiation of germ cells, gonad development and gametogenesis. In this study, PGC were isolated from Hailan chicken embryo, and then subcultured on feed layer with somatic cells after initial primary culture. In addition, the effect of RA on proliferation of cultured PGC and meiosis initiation of germ cells in embryonic chicken ovary were evaluated, together with underlining mechanisms. Meanwhile, we also investigate the production of transgenic chicken by LV-siRNA and get a model of Raldh2knock down. These studies will provide theoretic guidance for further investigation of early development and preparation of transgenic poultry.
1. Isolation, culture and identification of chicken PGC
PGC were collected from stage14chicken embryonic bloods, stage19genital ridges or stage28gonads with a fine glass needle under a microsurgery scope. For primary culture, cell suspension containing both PGC and somatic cells was seeded onto gelatin-treated6-well culture plates at a density of1×106/well in KO-DMEM supplemented with7.5%fetal calf serum (FCS),10ng/ml leukemia inhibitory factor (LIF),10ng/ml human basic fibroblast growth factor (bFGF),0.1mM MEM nonessential amino acids.0.1mM2-mercaptoethanol,2mM L-glutamine (Glu),100U/ml penicillin and100μg/ml streptomycin, PGC were cultured at38.5℃in an air atmosphere containing5%CO2with60%-70%relative humidity. To trace the origin of the colonies, the primary formed colonies were picked up with a fine glass needle and then subjected to RT-PCR analysis for expression of PGC-specific markers. For further subculture, colonies that were positive for PGC markers were picked up and digested to single cell suspension and reseeded onto6-well dish. The third passage PGC were identified by staining of Alkaline phosphatase (AKP), periodic acid-Schiff regent (PAS), c-kit, stage-specific embryonic antigens (SSEA-1,3.4) and EMA-1immunocytochemistry, and the expression of the pluripotency-associated genes PouV, Nanog and Sox2was analyzed by RT-PCR. The results all confirmed the characteristics of cultured PGC which indicated that the primary and subculture models of PGC could be used for studies about regulation of PGC proliferation and differenciation.
2. Effects of RA on adhesion and proliferation of chicken PGC
In the present study, the proliferating effect of RA on PGC was investigated along with the intracellular PI3K/Akt-mediated NF-κB signaling cascade and PKC/(3-catenin signaling. Results showed that RA significantly promoted PGC proliferation in a dose-and time-dependent manner, which was confirmed by BrdU incorporation. RA induced PGC aggregation by increasing expression of E-cadherin and α/β-catenins. However, this promoting effect was attenuated obviously by sequential inhibitors of LY294002for PI3K, KP372-1, for Akt SN50for NF-κB and PKC inhibitor H7, respectively. Furthermore, Western blot analysis manifested increased Akt phosphorylation (Ser ") of PGC after stimulation with RA, but this effect was abolished by LY294002or KP372-1. Meanwhile, treatment with RA increased expression of NF-κB and decreased IκBα expression that were inhibited by SN50. Moreover, blockade of PI3K or Akt activity resulted in inhibiting NF-κB translocation from the cytoplasm to the nucleus. E-cadherin and β-catenin protein expression levels also were increased by RA addition. Finally, the up-regulated mRNA expression of cell cycle regulating genes (cyclin D1and E, cyclin-dependent kinases6and2) was observed in the RA-treated cells. Flow cytometry analysis confirmed that RA-treated cultured PGC populations displayed a significant increase in the proportion of S and G2phase cells. Again, this stimulation was remarkably retarded by combined treatment with LY294002, KP372-1, SN50and H7respectively. Therefore, these results suggest that RA promote proliferation of the cultured chicken PGC via the PI3K/Akt mediated NF-κB signaling cascade and PKC/β-catenin signaling pathway.
3. Effect of RA on meiosis initiation in the chicken embryo
In the present study, the effect of RA on meiosis initiation in embryonic germ cells was investigated by organ culture in vitro and RNA interference. The results showed the profile of meiosis in chicken embryo ovary-meiotic germ cells were increased gradually since day15.5(first detectable) to day18.5(widespread) by immunofluorescence of SCP3and yH2AX. And the expression of Stra8is specifically up-regulated at day12.5, Scp3and Dmcl became elevated at day15.5. In addition, we observed increased mRNA levels for Raldh2but deceased gradually for Cyp26bl during meiosis that evidence for a requirement of RA accumulation to sustain meiosis. The expression of RARβ also presented slightly increase after day15.5. But in males, the expression of these genes keeps low level throughout development. We also found that the culture ovaries from day10.5and12.5were able to initiate meiosis, and meiosis is stimulated by RA compared with control (P<0.05). Finally, Raldh2shRNA remarkably abolished RA-dependent functions, indicating that RA synthesis and signaling is crucial for meiosis initiation. Taken together, these studies indicate that RA and signal are important in regulating the onset of meiosis in the chicken embryonic ovary. We reported here for the first time that Raldh2knock-down could significantly reduced germ cells to enter meiosis.
4. Production of transgenic chicken via LV-siRNA system
We designed three pairs of shRNA specific for Raldh2by RNA interference to knock down its function and constructed expressing vector of shRNA The day15.5ovarian cells were transfected with either shRNA specific for Raldh2. or a non-targeting shRNA as a negative control. The efficiency of transfection is about80%-90%. Among of these shRNAs, shRNA2produced the most significant inhibitory effect on Raldh2mRNA expression. RNA interference of Raldh2prevented the appearance of meiotic cells by immunofluorescence of SCP3. The Lentivirus vector specific for Raldh2shRNA was constructed and packaged. Then the virus was injected into X stage chicken embryos for production of transgenic chicken. The chicken model of Raldh2knock down was successfully obtained. The results showed that there were no obviously difference between transgenic chicken and wild chicken, as the morphology of ovary. The GFP was detected by PCR, which confirmed the extra gene was injected to genome successfully. The expression of Raldh2was decreased significantly by qRT-PCR. This research provide theoretical basis for understanding the molecular mechanisms of regulation of oogenesis and follicular development.
The above results indicated that the co-culture model of somatic cells and PGC from chicken embryo could be used to study on proliferation and differentiation. PGC were characterized by staining of AKP, PAS, c-kit, SSEA-1,3,4and EMA-1immunocytochemistry. The pluripotency of PGC was also detected by the expression of the pluripotency-associated genes (PouV,Nanog and Sox2). We investigated the molecular mechanisms of RA induced PGC proliferation by this model. Furthermore, we described the morphology and genetic changes during meiosis initiation in chicken embryonic ovary, then studied the regulating effect of RA on meiosis initiation by organ culture in vitro and RNA interference. These findings provide theoretic guidance and experimental platform for studying the development of germ cells and preparation of chimeras and transgenic poultry.
引文
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