肉鸭Myostatin剪切体的表达和调控
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摘要
肌肉生长抑制素(Myostatin, MSTN)是TGF-β (Transforming Growth Factor β)超家族的一员,对肌肉生长、发育具有负调控作用,其活性的丧失能引起动物肌纤维直径变大或肌纤维数增加,在哺乳动物中表现为“双肌”性状。所有脊椎动物中均存在MSTN基因,但其具有不同的基因组特征：鱼类由于DNA复制具有多个MSTN基因,而哺乳动物和鸟类仅有单拷贝的MSTN基因。大量的组织表达谱和发育表达谱研究显示,鱼类与哺乳动物MSTN基因表达谱具有物种差异,并极可能存在功能差异。最近研究发现禽类中MSTN基因存在多种剪切体。然而,目前对禽类MSTN剪切体的数量、剪切方式及表达模式尚缺乏研究。因此,本试验在前期扩增出MSTN基因剪切体EST序列的基础上,以北京鸭为研究材料,分别从mRNA和DNA水平研究了肉鸭MSTN的不同剪切体和剪切方式,同时利用荧光定量技术检测了各剪切体组织表达谱和发育表达谱,并进一步利用转染技术研究其在鸭胚成肌细胞中对下游基因的调控的作用。主要研究结果如下：
(1)利用RT-PCR、RACE等技术,扩增获得了北京鸭MSTN基因4种剪切变异体的mRNA序列,分别命名为MSTN-a、MSTN-b、MSTN-c和MSTN-d,同时证实了这4条mRNA序列具有相同的5'-UTR和3’-UTR序列。以所获得的鸭MSTN外显子序列为基础,通过PCR和克隆测序获得了该基因的内含子序列,进而拼接得到鸭MSTN基因部分DNA序列,长6,082bp。将4种剪切体序列与基因组DNA序列进行比对,发现北京鸭MSTN基因共有8个剪切位点,且各剪切位点均符合GU-AG剪切规则。北京鸭MSTN基因剪切体具有2种选择性剪接方式：外显子跳跃和3’剪切位点选择性使用,选择性剪切导致了剪切变异体结构与功能的改变。MSTN-b和MSTN-c由于提前终止子的出现,导致蛋白序列中RSRR结构和成熟区结构丢失,但其仍具有部分N-端LAP结构；MSTN-d却与MSTN-a相同,具有完整C-端成熟区结构和RSRR结构。
(2)通过荧光定量技术检测了MSTN基因4种剪切体在30d(天)北京鸭13个组织中的表达水平,发现各剪切体的表达均具有组织特异性和性别差异性。
(3)通过荧光定量技术检测了MSTN基因4种剪切体在北京鸭胸肌、腿肌和心肌中的发育表达模式,结果表明：MSTN-a在胸肌和腿肌中的发育表达差异不显著,但表达水平较高,并且MSTN-a在胚胎心脏发育后期表达降低,可能对心脏发育起到了一定的调控作用；MSTN-b在腿肌中的表达水平高于胸肌,推测它可能主要是在腿肌中发挥作用；MSTN-c在胸肌和腿肌中的表达量均处于极低的水平,可能不参与调控肌肉的发育过程；MSTN-d在北京鸭腿肌和胸肌中的表达水平虽然很低,但具有完全不同的表达模式,推测可能与肌纤维类型有关。
(4)转染MSTN-a和MSTN-b到鸭胚成肌细胞后,发现转染MSTN-a组的MyoD表达量有显著的下调；而转染MSTN-b之后,MyoG和Akirin2的表达有所下调,推测在鸭中,MSTN-a可能参与了成肌细胞增殖过程的调控,而MSTN-b可能与成肌细胞分化过程有关。
MSTN (Myostatin) is a member of transforming growth fator-β superfamily (TGF-β) and plays a negative regulatory role in skeletal muscle growth and development. The inactive MSTN can give rise to diameter and number of muscle fiber increasing significantly, and called "double muscling" in mammals. The MSTN gene was found in vertebrates, and has different genome characters:single copy gene in mammals and birds and four MSTN genes in fish causing by gene duplication. Tissue-specific and developing expression profiles showed different expression patterns between fish and mammals, suggesting functional diverged in MSTN genes. Recently, alternative splicing isoform of MSTN genes were found in bird, however, there were no further reports about the numbers, splicing sites and expression profiles of alternative splicing isoform of MSTN in duck. Based on ESTs of duck MSTN, which were obtained by preliminary tests, we employed Peking duck as materials to amplify the mRNA and DNA sequence of duck MSTN sequences, and to analyze the alternative splicing sites, tissue-specific and developing expression profiles, and then alternative splicing isoform transfection assay was performed to detect the regulation roles to downstream gene expression. The main results of this study are as follows:
(1) RT-PCR and RACE assays were performed to obtain the full-mRNA and DNA sequences of alternative splicing isoform of duck MSTN gene. Finally, we have got a DNA sequences with length6,082bp, and four splicing isoforms of duck MSTN gene and were named MSTN-a, MSTN-b, MSTN-c and MSTN-d. Based on the comparison between each mRNA variant and the DNA sequence, we found that all the intron splicing of duck MSTN were consistent with GU-AG rules with8splicing sites, and within two alternative splicing methods:exon skipping and3'SSs selective using. The structure and function were changed by alternative splicing:Because of premature stop codon and the deletion of RSRR site and mature domain, the MSTN-b and MSTN-c retained only the N-terminal LAP domains. However, MSTN-d was not missing these domains, in contrast to MSTN-a.
(2) The qRT-PCR assay was performed to analysis the expression profiles in30d Peking duck with13tissues and found that the alternative splicing isoform of duck MSTN has different expression profiles between tissues and sex.
(3) The real-time PCR results showed that MSTN-a was expressed in breast muscle, leg muscle and heart with higher level than MSTN-b, MSTN-c and MSTN-d. There was no significant difference between breast muscle and leg muscle in MSTN-a mRNA expression, and MSTN-a was found to have essential roles during heart development at late stage. MSTN-b was thought to play roles during leg muscle development with higher expression level than which in breast muscle. MSTN-c has significant lower expression levels in breast muscle and leg muscle, and could not have function in regulating muscle development. During duck embryo muscle development, MSTN-d showed the different expression patterns between leg muscle and breast muscle, suggesting that it should relate to muscle fiber type.
(4) The transfection assay showed that, MyoD was down-regulated by MSTN-a, and Akirin2and MyoG were down-regulated by MSTN-b, showing that MSTN-a probably have major roles in myoblast proliferation, and MSTN-b in differentiation.
(1)利用RT-PCR、RACE等技术,扩增获得了北京鸭MSTN基因4种剪切变异体的mRNA序列,分别命名为MSTN-a、MSTN-b、MSTN-c和MSTN-d,同时证实了这4条mRNA序列具有相同的5'-UTR和3’-UTR序列。以所获得的鸭MSTN外显子序列为基础,通过PCR和克隆测序获得了该基因的内含子序列,进而拼接得到鸭MSTN基因部分DNA序列,长6,082bp。将4种剪切体序列与基因组DNA序列进行比对,发现北京鸭MSTN基因共有8个剪切位点,且各剪切位点均符合GU-AG剪切规则。北京鸭MSTN基因剪切体具有2种选择性剪接方式：外显子跳跃和3’剪切位点选择性使用,选择性剪切导致了剪切变异体结构与功能的改变。MSTN-b和MSTN-c由于提前终止子的出现,导致蛋白序列中RSRR结构和成熟区结构丢失,但其仍具有部分N-端LAP结构；MSTN-d却与MSTN-a相同,具有完整C-端成熟区结构和RSRR结构。
(2)通过荧光定量技术检测了MSTN基因4种剪切体在30d(天)北京鸭13个组织中的表达水平,发现各剪切体的表达均具有组织特异性和性别差异性。
(3)通过荧光定量技术检测了MSTN基因4种剪切体在北京鸭胸肌、腿肌和心肌中的发育表达模式,结果表明：MSTN-a在胸肌和腿肌中的发育表达差异不显著,但表达水平较高,并且MSTN-a在胚胎心脏发育后期表达降低,可能对心脏发育起到了一定的调控作用；MSTN-b在腿肌中的表达水平高于胸肌,推测它可能主要是在腿肌中发挥作用；MSTN-c在胸肌和腿肌中的表达量均处于极低的水平,可能不参与调控肌肉的发育过程；MSTN-d在北京鸭腿肌和胸肌中的表达水平虽然很低,但具有完全不同的表达模式,推测可能与肌纤维类型有关。
(4)转染MSTN-a和MSTN-b到鸭胚成肌细胞后,发现转染MSTN-a组的MyoD表达量有显著的下调；而转染MSTN-b之后,MyoG和Akirin2的表达有所下调,推测在鸭中,MSTN-a可能参与了成肌细胞增殖过程的调控,而MSTN-b可能与成肌细胞分化过程有关。
MSTN (Myostatin) is a member of transforming growth fator-β superfamily (TGF-β) and plays a negative regulatory role in skeletal muscle growth and development. The inactive MSTN can give rise to diameter and number of muscle fiber increasing significantly, and called "double muscling" in mammals. The MSTN gene was found in vertebrates, and has different genome characters:single copy gene in mammals and birds and four MSTN genes in fish causing by gene duplication. Tissue-specific and developing expression profiles showed different expression patterns between fish and mammals, suggesting functional diverged in MSTN genes. Recently, alternative splicing isoform of MSTN genes were found in bird, however, there were no further reports about the numbers, splicing sites and expression profiles of alternative splicing isoform of MSTN in duck. Based on ESTs of duck MSTN, which were obtained by preliminary tests, we employed Peking duck as materials to amplify the mRNA and DNA sequence of duck MSTN sequences, and to analyze the alternative splicing sites, tissue-specific and developing expression profiles, and then alternative splicing isoform transfection assay was performed to detect the regulation roles to downstream gene expression. The main results of this study are as follows:
(1) RT-PCR and RACE assays were performed to obtain the full-mRNA and DNA sequences of alternative splicing isoform of duck MSTN gene. Finally, we have got a DNA sequences with length6,082bp, and four splicing isoforms of duck MSTN gene and were named MSTN-a, MSTN-b, MSTN-c and MSTN-d. Based on the comparison between each mRNA variant and the DNA sequence, we found that all the intron splicing of duck MSTN were consistent with GU-AG rules with8splicing sites, and within two alternative splicing methods:exon skipping and3'SSs selective using. The structure and function were changed by alternative splicing:Because of premature stop codon and the deletion of RSRR site and mature domain, the MSTN-b and MSTN-c retained only the N-terminal LAP domains. However, MSTN-d was not missing these domains, in contrast to MSTN-a.
(2) The qRT-PCR assay was performed to analysis the expression profiles in30d Peking duck with13tissues and found that the alternative splicing isoform of duck MSTN has different expression profiles between tissues and sex.
(3) The real-time PCR results showed that MSTN-a was expressed in breast muscle, leg muscle and heart with higher level than MSTN-b, MSTN-c and MSTN-d. There was no significant difference between breast muscle and leg muscle in MSTN-a mRNA expression, and MSTN-a was found to have essential roles during heart development at late stage. MSTN-b was thought to play roles during leg muscle development with higher expression level than which in breast muscle. MSTN-c has significant lower expression levels in breast muscle and leg muscle, and could not have function in regulating muscle development. During duck embryo muscle development, MSTN-d showed the different expression patterns between leg muscle and breast muscle, suggesting that it should relate to muscle fiber type.
(4) The transfection assay showed that, MyoD was down-regulated by MSTN-a, and Akirin2and MyoG were down-regulated by MSTN-b, showing that MSTN-a probably have major roles in myoblast proliferation, and MSTN-b in differentiation.
引文
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