Shugoshin1对牛卵母细胞成熟、早期胚胎发育及体细胞有丝分裂影响的研究
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摘要
有丝分裂和减数分裂当中染色体分离具有严格的时空性,使细胞如实的将遗传信息传递给后代。姐妹染色单体连接在染色体分离中扮演着重要的角色,这种连接的主要中介是环状多蛋白复合物cohesirin。
Shugoshin(Sgo/SGO)在果蝇、玉米、水稻、酵母、非洲爪蟾、小鼠及Hela细胞染色体分离当中的作用都有研究报道,该蛋白家族的主要作用是保护着丝粒粘连蛋白cohesin,维持姐妹染色单体的连接,特别是在第一次减数分裂过程中。除此之外,Sgo/SGO在有丝分裂中还有多个作用,对于细胞分裂进程和染色体分离至关重要。目前对SGO在哺乳动物的正常成体细胞及卵母细胞中的研究及其少见,仅在小鼠中进行了有限的探索。
本研究分析了SGO1/Sgol在牛、大鼠和小鼠组织中的表达水平的变化,对牛体细胞中SGO1的功能进行了初步分析,确定了它在牛卵母细胞减数分裂过程中的分布位置,并进一步用超表达和低表达的方法探讨了SGO1在牛卵母细胞减数分离和早期胚胎发育过程中的作用。实验成果对于明晰脊椎动物特别是哺乳动物中SGO1的作用具有重要的参考价值,主要研究结果如下：
1.SGO1/Sgol在牛、大鼠和小鼠组织中的表达水平
通过实时定量PCR方法,检测了三个哺乳动物物种中SGO1/Sgol的mRNA水平。SGO1/Sgol的表达存在物种差异性、组织差异性和性别差异性,但其表达水平有一定的规律可循。SGO1/Sgol在大鼠和小鼠的胰腺、大脑、肾脏、心脏、肝脏、肌肉、延髓和肺等组织中的表达量都相对较低,而在小鼠、雄性大鼠和母牛脾脏中的表达水平较高,SGO1/Sgol在雌性大鼠小肠中的表达量最多,在公牛睾丸组织中高表达。
2.SGO1在牛胎儿成纤维细胞中的作用
通过向同步化的细胞中转染siRNA的方法来研究SGO1对哺乳动物细胞有丝分裂染色体分离的影响。削减SGO1的表达会引起着丝粒和染色体臂上的姐妹染色单体连接提前消失,造成牛胎儿成纤维细胞的有丝分裂阻滞。成对排列的无连接染色体无法移向细胞的两极,逐渐的凝集成短棒状,最后散在的单个染色体杂乱的分布于细胞中,形成多倍体细胞。
3.SGO1在牛卵母细胞减数分裂成熟过程中的定位
通过注射带有标记的外源SGO1mRNA到GV期卵母细胞胞质的方法,来分析SGO1在牛卵母细胞减数分裂染色体上的分布情况。SGO1自第一次减数分裂前中期至第二次减数分裂中期,都定位在染色体上,但并不完全重合,它并不分布于整条染色体臂上,而是在某些部位呈现点状富集。在第一次减数分裂中,每条染色体上都有SGO1信号,进入第二次减数分裂以后,SGO1只在核上有表达并在某些位点显示较强的着色,极体中检测不到SGO1信号。
4.SGO1对牛卵母细胞减数分裂和早期胚胎发育的影响
通过显微注射SGO1特异的siRNA或者mRNA到卵母细胞胞质的方法,探讨超表达和低表达SGO1对卵母细胞分裂进程和植入前胚胎发育的影响。SGO1参与调控染色体的分离。外源过表达SGO1不会影响同源染色体的分离,但分开的染色体无法移向细胞的两极,呈现一个不规则的中期样排列。干扰SGO1的表达会引起卵母细胞分裂阻滞,早期胚胎发育受到影响,囊胚率和囊胚细胞数减少。
The chromosomal separation in both mitosis and meiosis is time-dependent and space-dependent; ensuring the faithful delivery of genetic materials.Sister chromatid cohesion plays an important role in chromosomal separation, which is mediated by cohesin, a circular multi-subunit protein complex.
It is reported that Shugoshin(Sgo/SGO) involves in chromosomal separation in fruit fly,maize,rice,yeast,xenopus,mouse and Hela cells.The major role of such protein family is to protect cohesin at centromere and maintain the cohesion between sister chromatids,especially in meiosis I.Besides,Sgo/SGO functions in several other process and is essential for cell division and chromosomal separation.It is hard to see the relevant researches about mammalian Sgo/SGO in normal somatic cells and oocytes, if any, just in mouse.
In this research, the amount of SGO1/Sgo1mRNA in different tissues of bovine, rat and mouse was detected firstly. The roles of SGO1was proved in bovine somatic cells subsequently.We also confirmed the distribution of SGO1during oocytes meiotic maturation.The roles of SGO1during bovine oocyte meiosis and early embryo development were further discussed via its over expression and low expression. The results as follows provide an important reference for researches in vertebrate animals, especially for mammals.
1. The expression levels of SGO1/Sgol in bovine, rat and mouse tissues
The mRNA levels of SGO1/Sgol in three species were detected by real time RT PCR. The expression of SGO1/Sgol was different among species, tissues and genders, but regularities were drawed The expression levels of SGO1/Sgol were lower in pancreas, brain, kidney, heart, liver, muscle, medulla oblongata and lung of rat and mouse, but higher in spleen of mouse, male rat and female bovine, as well as the most detection in small intestine of female rat and testis of male bovine.
2. Roles of SGO1in bovine embryonic fibroblasts
siRNA was transfected to the synchronized cells to clarify the function of SGO1in mitosis of mammalian cells.Depletion of SGO1promoted the premature loss of sister chromatid cohesion along chromosome arms and at centromeres and caused mitotic arrestment.The chromosome aligned in pair although with no cohesion.Then they hypercondensed and turned to short and thick rods dispersed disorderly without being pulled to the opposite spindle poles. Such polyploid cells failed to enter anaphase increased over time.
3. Localization of SGO1during bovine oocyte meiotic maturation
The localization of SGO1on meiotic chromosomes was analysed by injection of marked SGO1mRNA to the cytoplasm of GV oocytes. SGO1anchored on chromosomes from prometaphase I to metaphase II, however, not lapped entirely, with several enrichment spots. SGO1signals were detected on every chromosome in meiosis Ⅰ, but appeared only at nucleus and presented some strong staining spots when entry to meiosis Ⅱ, not detected in polar body.
4. Effect of SGO1on bovine oocyte meiosis and early embryo development
SGO specific siRNA or mRNA was microinjected to oocyte cytoplasm in order to explore the influence of over expression and depletion of SGO1on oocyte meiotic division and early embryo development. SGOl was involved in chromosomal separation. The homologous chromosomes could separate apart from each other after exogenous overexpression of SGO1but not be pulled to the opposite poles of the spindle, resulting a metaphase-like appearance. Depletion of SGO1at GV stage caused oocyte division arrestment, lower blastocyst rate and blastocyst with fewer cells.
Shugoshin(Sgo/SGO)在果蝇、玉米、水稻、酵母、非洲爪蟾、小鼠及Hela细胞染色体分离当中的作用都有研究报道,该蛋白家族的主要作用是保护着丝粒粘连蛋白cohesin,维持姐妹染色单体的连接,特别是在第一次减数分裂过程中。除此之外,Sgo/SGO在有丝分裂中还有多个作用,对于细胞分裂进程和染色体分离至关重要。目前对SGO在哺乳动物的正常成体细胞及卵母细胞中的研究及其少见,仅在小鼠中进行了有限的探索。
本研究分析了SGO1/Sgol在牛、大鼠和小鼠组织中的表达水平的变化,对牛体细胞中SGO1的功能进行了初步分析,确定了它在牛卵母细胞减数分裂过程中的分布位置,并进一步用超表达和低表达的方法探讨了SGO1在牛卵母细胞减数分离和早期胚胎发育过程中的作用。实验成果对于明晰脊椎动物特别是哺乳动物中SGO1的作用具有重要的参考价值,主要研究结果如下：
1.SGO1/Sgol在牛、大鼠和小鼠组织中的表达水平
通过实时定量PCR方法,检测了三个哺乳动物物种中SGO1/Sgol的mRNA水平。SGO1/Sgol的表达存在物种差异性、组织差异性和性别差异性,但其表达水平有一定的规律可循。SGO1/Sgol在大鼠和小鼠的胰腺、大脑、肾脏、心脏、肝脏、肌肉、延髓和肺等组织中的表达量都相对较低,而在小鼠、雄性大鼠和母牛脾脏中的表达水平较高,SGO1/Sgol在雌性大鼠小肠中的表达量最多,在公牛睾丸组织中高表达。
2.SGO1在牛胎儿成纤维细胞中的作用
通过向同步化的细胞中转染siRNA的方法来研究SGO1对哺乳动物细胞有丝分裂染色体分离的影响。削减SGO1的表达会引起着丝粒和染色体臂上的姐妹染色单体连接提前消失,造成牛胎儿成纤维细胞的有丝分裂阻滞。成对排列的无连接染色体无法移向细胞的两极,逐渐的凝集成短棒状,最后散在的单个染色体杂乱的分布于细胞中,形成多倍体细胞。
3.SGO1在牛卵母细胞减数分裂成熟过程中的定位
通过注射带有标记的外源SGO1mRNA到GV期卵母细胞胞质的方法,来分析SGO1在牛卵母细胞减数分裂染色体上的分布情况。SGO1自第一次减数分裂前中期至第二次减数分裂中期,都定位在染色体上,但并不完全重合,它并不分布于整条染色体臂上,而是在某些部位呈现点状富集。在第一次减数分裂中,每条染色体上都有SGO1信号,进入第二次减数分裂以后,SGO1只在核上有表达并在某些位点显示较强的着色,极体中检测不到SGO1信号。
4.SGO1对牛卵母细胞减数分裂和早期胚胎发育的影响
通过显微注射SGO1特异的siRNA或者mRNA到卵母细胞胞质的方法,探讨超表达和低表达SGO1对卵母细胞分裂进程和植入前胚胎发育的影响。SGO1参与调控染色体的分离。外源过表达SGO1不会影响同源染色体的分离,但分开的染色体无法移向细胞的两极,呈现一个不规则的中期样排列。干扰SGO1的表达会引起卵母细胞分裂阻滞,早期胚胎发育受到影响,囊胚率和囊胚细胞数减少。
The chromosomal separation in both mitosis and meiosis is time-dependent and space-dependent; ensuring the faithful delivery of genetic materials.Sister chromatid cohesion plays an important role in chromosomal separation, which is mediated by cohesin, a circular multi-subunit protein complex.
It is reported that Shugoshin(Sgo/SGO) involves in chromosomal separation in fruit fly,maize,rice,yeast,xenopus,mouse and Hela cells.The major role of such protein family is to protect cohesin at centromere and maintain the cohesion between sister chromatids,especially in meiosis I.Besides,Sgo/SGO functions in several other process and is essential for cell division and chromosomal separation.It is hard to see the relevant researches about mammalian Sgo/SGO in normal somatic cells and oocytes, if any, just in mouse.
In this research, the amount of SGO1/Sgo1mRNA in different tissues of bovine, rat and mouse was detected firstly. The roles of SGO1was proved in bovine somatic cells subsequently.We also confirmed the distribution of SGO1during oocytes meiotic maturation.The roles of SGO1during bovine oocyte meiosis and early embryo development were further discussed via its over expression and low expression. The results as follows provide an important reference for researches in vertebrate animals, especially for mammals.
1. The expression levels of SGO1/Sgol in bovine, rat and mouse tissues
The mRNA levels of SGO1/Sgol in three species were detected by real time RT PCR. The expression of SGO1/Sgol was different among species, tissues and genders, but regularities were drawed The expression levels of SGO1/Sgol were lower in pancreas, brain, kidney, heart, liver, muscle, medulla oblongata and lung of rat and mouse, but higher in spleen of mouse, male rat and female bovine, as well as the most detection in small intestine of female rat and testis of male bovine.
2. Roles of SGO1in bovine embryonic fibroblasts
siRNA was transfected to the synchronized cells to clarify the function of SGO1in mitosis of mammalian cells.Depletion of SGO1promoted the premature loss of sister chromatid cohesion along chromosome arms and at centromeres and caused mitotic arrestment.The chromosome aligned in pair although with no cohesion.Then they hypercondensed and turned to short and thick rods dispersed disorderly without being pulled to the opposite spindle poles. Such polyploid cells failed to enter anaphase increased over time.
3. Localization of SGO1during bovine oocyte meiotic maturation
The localization of SGO1on meiotic chromosomes was analysed by injection of marked SGO1mRNA to the cytoplasm of GV oocytes. SGO1anchored on chromosomes from prometaphase I to metaphase II, however, not lapped entirely, with several enrichment spots. SGO1signals were detected on every chromosome in meiosis Ⅰ, but appeared only at nucleus and presented some strong staining spots when entry to meiosis Ⅱ, not detected in polar body.
4. Effect of SGO1on bovine oocyte meiosis and early embryo development
SGO specific siRNA or mRNA was microinjected to oocyte cytoplasm in order to explore the influence of over expression and depletion of SGO1on oocyte meiotic division and early embryo development. SGOl was involved in chromosomal separation. The homologous chromosomes could separate apart from each other after exogenous overexpression of SGO1but not be pulled to the opposite poles of the spindle, resulting a metaphase-like appearance. Depletion of SGO1at GV stage caused oocyte division arrestment, lower blastocyst rate and blastocyst with fewer cells.
引文
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