斑马鱼secl3基因在器官发育过程中的功能研究
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摘要
作为一种越来越受关注的模式生物,斑马鱼因其自身所具有的优势已在生命科学的研究中发挥着越来越大的作用。斑马鱼早期胚胎发育是一个由多基因和信号通路共同调控的过程。任何一个参与这一调控过程的基因或信号通路缺陷都会导致早期胚胎发育异常。
本研究主要利用遗传学,细胞生物学以及分子生物学的研究方法研究sec13基因在斑马鱼早期胚胎发育中的作用。通过ENU介导的突变筛选,我们实验室获得了一个sec13突变的斑马鱼品系secl3sq198。我们前期的研究显示在see13sq198突变体中sec13发生了一个T→A的点突变,该点突变发生在sec13基因第七个内含子倒数第十个核苷酸。这个点突变导致sec13基因的mRNA前体hnRNA在发生剪切的时候剪切位点前移,最终产生了多了8个核苷酸的异常的mRNA。这一异常的mRNA在翻译Sec13蛋白的时候出现了阅读框移码,导致蛋白产物的C端缺失了正常Sec13的85个氨基酸,同时又增加了34个新的氨基酸。而且我们前期的实验还证明该C端缩短的Sec13已失去了与其互作蛋白Sec31a互作的能力。在此基础上我们深入地研究了Sec13在斑马鱼早期器官发育中的功能。
在真核细胞内,Sec13有两个功能：一是与Sec31相互作用形成COPII复合体的外围复合物参与内质网到高尔基体的蛋白转运；二是与Nup145C(酵母)或Nup96(脊椎动物)以及与其他核孔蛋白相互作用参与核孔复合物的形成从而调节核质转运和基因表达调控。我们第一部分的工作主要是研究作为COPⅡ复合体组成部分的Sec13在斑马鱼早期肠道及其附属器官(肝脏,胰腺)发育中的作用。我们的研究结果发现,C端缩短的Secl3蛋白失去了与Sec31a互作的能力从而导致COPⅡ功能受损。结果,COPII功能受损导致内质网到高尔基体的蛋白转运受阻,从而使细胞外分泌蛋白大量积累在软骨细胞,肝脏细胞和肠道细胞的内质网中。内质网中蛋白过度积累使这些细胞出现了非折叠蛋白反应。最终突变体斑马鱼肠道,肝脏和胰腺由于细胞周期受阻和细胞凋亡而表现出小肠道,小肝脏和小胰腺的表型。
我们第二部分的工作主要是研究作为核孔复合体组成部分的Sec13在斑马鱼早期视网膜发育中的作用。我们的研究发现,sec13sq198突变体不仅表现出变小的视网膜,而且其内部的分层结构严重遭到破坏(感受器细胞层几乎完全消失,其余两层细胞数量大大减少)。进一步的研究发现,sec13sq198突变体视网膜细胞的分化是相对正常的。最重要的是,我们发现secl3sq198突变体中破坏的视网膜结构不是由COPⅡ功能受损引起的,因为当我们把COPⅡ复合体的另两个组分Sec31a和Sec31b同时敲除或者用药物阻断内质网到高尔基体的蛋白转运时,斑马鱼胚胎眼睛虽然变小,但其内部的分层结构并未受到破坏。之后,当我们通过电子显微镜和免疫荧光观察sec13sq198突变体视网膜不同类型细胞的核孔结构时我们发现这些核孔的形成严重受到阻碍。而且,我们发现核孔结构形成异常导致总1nRNA累积在视网膜不同类型的细胞核内。最后突变体斑马鱼由于激活的p53所介导的视网膜细胞凋亡而表现出小眼睛表型。
综上所述,利用sec13sq198这个遗传突变体,我们不仅证明了Sec13在消化器官和视网膜发育中起重要作用,而且还首次将Sec13的两个功能在多细胞生物水平上很好的区分开了。
As a model system which is receiving increasing attentions, zebrafish is playing a growing role in life sciences research. Early embryogenesis of zebrafish is a complex processes controlled by a large number of genes and different signaling pathways. Any defects in these processes can lead to embryonic abnormalities.
In this study, we utilized genetic, cell biology and molecular biology approaches to investigate the role of sec13gene during zebrafish early embryogenesis. Through ENU-mediated mutagenesis, we obtained a zebrafish sec13mutant, sec13sq198. Our previous study showed that there is a T to A transversion in the7th intron of the sec13gene. This point mutation creates a new splicing receptor site which leads to the addition of extra eight nucleotides into the final mRNA. This mutant mRNA is predicted to encode a polypeptide with85amino acids truncated from the carboxyl-terminus of WT Secl3and the addition of34new amino acids due to frame shift in the new open reading frame. Furthermore, Our previous work also demonstrated that carboxyl-terminus-truncated Sec13loses its affinity to Sec31a. So based on these data, we decide to comprehensively investigate the role of sec13during zebrafish early embryogenesis.
Sec13is a dual functional protein in eukaryotes. First, it interacts with Sec31a to form the outer coat of the COPII complex, facilitating protein transport form ER to Golgi apparatus. Second, it also interacts with (Nup145C in yeast and Nup96in vertebrates) to form NPCs to facilitate nucleo-cytoplasmic transport and regulate gene expression. Our first part efforts mainly focused on the role of sec13during zebrafish digestive organs organogenesis, namely, the intestine, liver and pancreas. We found that the carboxyl-terminus-truncated Sec13in secl3sq198loses its affinity to Sec31a, that leads to the dysfucntuion of the COPII complex. As a consequence, protein transport from ER to golgi apparatus was blocked, giving rise to the accumulation of secretory protein in the ER of chondrocytes, liver and intestinal cells. Unfolded protein response (UPR) was activated due to the sudden build up of protein mass in ER and consequently, the stressed cells underwent cell-cycle arrest and cell apoptosis, which halt the growth of developing digestive organs.
Our second part is about nuclear pore function of sec13during retinal early development. In this respect, we found that, despite that all types of the retinal cells were specified, sec13sq198mutant embryos developed small eyes characterized by gradual loss of the photoreceptor cell layer (PCL) and reduction of cell numbers in other cell layers. Surprisingly, blocking COPII function either by double-knockdown of Sec31a and Sec31b or by drug treatment (Brefeldin A), though produced small eyes, did not disrupt the layered structure of the eye, suggesting that the retina lesion observed in sec13sq198was not totally due to COPII dysfunction. Our further data from immunostaining and transmission electron microscopy (TEM) analysis demonstrated that the retina of sec13sq198failed to form proper nuclear pores which led to the nuclear accumulation of total mRNA. TUNEL assay revealed that the p53-dependent apoptosis pathway was abnormally activated which finally caused the small eye phenotype by triggering cell apoptosis in sec13sq198.
Our overall data not only demonstrated the importantce of Sec13in organogenesis of digestive organs and eyes but also provided the first genetic evidence to demonstrate that the two functions of Sec13, namely as a key component of COPII and NPCs can be differentially uncoupled during the process of organogenesis in a multicellular organism.
本研究主要利用遗传学,细胞生物学以及分子生物学的研究方法研究sec13基因在斑马鱼早期胚胎发育中的作用。通过ENU介导的突变筛选,我们实验室获得了一个sec13突变的斑马鱼品系secl3sq198。我们前期的研究显示在see13sq198突变体中sec13发生了一个T→A的点突变,该点突变发生在sec13基因第七个内含子倒数第十个核苷酸。这个点突变导致sec13基因的mRNA前体hnRNA在发生剪切的时候剪切位点前移,最终产生了多了8个核苷酸的异常的mRNA。这一异常的mRNA在翻译Sec13蛋白的时候出现了阅读框移码,导致蛋白产物的C端缺失了正常Sec13的85个氨基酸,同时又增加了34个新的氨基酸。而且我们前期的实验还证明该C端缩短的Sec13已失去了与其互作蛋白Sec31a互作的能力。在此基础上我们深入地研究了Sec13在斑马鱼早期器官发育中的功能。
在真核细胞内,Sec13有两个功能：一是与Sec31相互作用形成COPII复合体的外围复合物参与内质网到高尔基体的蛋白转运；二是与Nup145C(酵母)或Nup96(脊椎动物)以及与其他核孔蛋白相互作用参与核孔复合物的形成从而调节核质转运和基因表达调控。我们第一部分的工作主要是研究作为COPⅡ复合体组成部分的Sec13在斑马鱼早期肠道及其附属器官(肝脏,胰腺)发育中的作用。我们的研究结果发现,C端缩短的Secl3蛋白失去了与Sec31a互作的能力从而导致COPⅡ功能受损。结果,COPII功能受损导致内质网到高尔基体的蛋白转运受阻,从而使细胞外分泌蛋白大量积累在软骨细胞,肝脏细胞和肠道细胞的内质网中。内质网中蛋白过度积累使这些细胞出现了非折叠蛋白反应。最终突变体斑马鱼肠道,肝脏和胰腺由于细胞周期受阻和细胞凋亡而表现出小肠道,小肝脏和小胰腺的表型。
我们第二部分的工作主要是研究作为核孔复合体组成部分的Sec13在斑马鱼早期视网膜发育中的作用。我们的研究发现,sec13sq198突变体不仅表现出变小的视网膜,而且其内部的分层结构严重遭到破坏(感受器细胞层几乎完全消失,其余两层细胞数量大大减少)。进一步的研究发现,sec13sq198突变体视网膜细胞的分化是相对正常的。最重要的是,我们发现secl3sq198突变体中破坏的视网膜结构不是由COPⅡ功能受损引起的,因为当我们把COPⅡ复合体的另两个组分Sec31a和Sec31b同时敲除或者用药物阻断内质网到高尔基体的蛋白转运时,斑马鱼胚胎眼睛虽然变小,但其内部的分层结构并未受到破坏。之后,当我们通过电子显微镜和免疫荧光观察sec13sq198突变体视网膜不同类型细胞的核孔结构时我们发现这些核孔的形成严重受到阻碍。而且,我们发现核孔结构形成异常导致总1nRNA累积在视网膜不同类型的细胞核内。最后突变体斑马鱼由于激活的p53所介导的视网膜细胞凋亡而表现出小眼睛表型。
综上所述,利用sec13sq198这个遗传突变体,我们不仅证明了Sec13在消化器官和视网膜发育中起重要作用,而且还首次将Sec13的两个功能在多细胞生物水平上很好的区分开了。
As a model system which is receiving increasing attentions, zebrafish is playing a growing role in life sciences research. Early embryogenesis of zebrafish is a complex processes controlled by a large number of genes and different signaling pathways. Any defects in these processes can lead to embryonic abnormalities.
In this study, we utilized genetic, cell biology and molecular biology approaches to investigate the role of sec13gene during zebrafish early embryogenesis. Through ENU-mediated mutagenesis, we obtained a zebrafish sec13mutant, sec13sq198. Our previous study showed that there is a T to A transversion in the7th intron of the sec13gene. This point mutation creates a new splicing receptor site which leads to the addition of extra eight nucleotides into the final mRNA. This mutant mRNA is predicted to encode a polypeptide with85amino acids truncated from the carboxyl-terminus of WT Secl3and the addition of34new amino acids due to frame shift in the new open reading frame. Furthermore, Our previous work also demonstrated that carboxyl-terminus-truncated Sec13loses its affinity to Sec31a. So based on these data, we decide to comprehensively investigate the role of sec13during zebrafish early embryogenesis.
Sec13is a dual functional protein in eukaryotes. First, it interacts with Sec31a to form the outer coat of the COPII complex, facilitating protein transport form ER to Golgi apparatus. Second, it also interacts with (Nup145C in yeast and Nup96in vertebrates) to form NPCs to facilitate nucleo-cytoplasmic transport and regulate gene expression. Our first part efforts mainly focused on the role of sec13during zebrafish digestive organs organogenesis, namely, the intestine, liver and pancreas. We found that the carboxyl-terminus-truncated Sec13in secl3sq198loses its affinity to Sec31a, that leads to the dysfucntuion of the COPII complex. As a consequence, protein transport from ER to golgi apparatus was blocked, giving rise to the accumulation of secretory protein in the ER of chondrocytes, liver and intestinal cells. Unfolded protein response (UPR) was activated due to the sudden build up of protein mass in ER and consequently, the stressed cells underwent cell-cycle arrest and cell apoptosis, which halt the growth of developing digestive organs.
Our second part is about nuclear pore function of sec13during retinal early development. In this respect, we found that, despite that all types of the retinal cells were specified, sec13sq198mutant embryos developed small eyes characterized by gradual loss of the photoreceptor cell layer (PCL) and reduction of cell numbers in other cell layers. Surprisingly, blocking COPII function either by double-knockdown of Sec31a and Sec31b or by drug treatment (Brefeldin A), though produced small eyes, did not disrupt the layered structure of the eye, suggesting that the retina lesion observed in sec13sq198was not totally due to COPII dysfunction. Our further data from immunostaining and transmission electron microscopy (TEM) analysis demonstrated that the retina of sec13sq198failed to form proper nuclear pores which led to the nuclear accumulation of total mRNA. TUNEL assay revealed that the p53-dependent apoptosis pathway was abnormally activated which finally caused the small eye phenotype by triggering cell apoptosis in sec13sq198.
Our overall data not only demonstrated the importantce of Sec13in organogenesis of digestive organs and eyes but also provided the first genetic evidence to demonstrate that the two functions of Sec13, namely as a key component of COPII and NPCs can be differentially uncoupled during the process of organogenesis in a multicellular organism.
引文
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