小麦雄性不育相关基因SKP1的克隆与表达分析
摘要
小麦(Triticum aestivum)是世界上重要的粮食作物,是人类得以生存繁衍的重要物质基础,与其它作物一样,利用杂种优势也是提高其产量和改良品质的最有效方法。然而,小麦杂种优势理论与实践研究还远不如玉米、水稻、油菜等作物那样广泛与深入。尤其是小麦雄性不育最佳利用途径与机理研究目前已成为攻克小麦杂种优势利用这一世界性难题的重要突破口。概括国内外小麦不育类型,主流研究可分为两大方面,即生理型雄性不育和遗传型雄性不育。对其研究,不同学者侧重点不同,从育性基因、育性差异蛋白、育性生理生化代谢途径等都进行了系列研究。泛素蛋白酶途径是细胞内冗余蛋白质降解的重要途径,其帮助细胞内冗余蛋白质降解,并能保证细胞内部复杂的生理生化过程正常进行。一类参与泛素蛋白酶途径的小分子量蛋白SKP1(S-phase kinase-associated-protein 1)广泛存在于真核生物中。其主要参与SCF复合体的形成,为维护SCF复合体的生物学功能起到了举足轻重的作用。SCF与细胞周期调控有关,其在生物体内主要是调控细胞内泛素介导的蛋白质降解过程,同时参与细胞内发育的多项生理过程。本研究正是基于SKP1上述作用,以化学杀雄剂SQ-1作为小麦雄性不育诱导剂,以小麦品种西农2611为试验材料,分喷药和不喷药(对照)两种处理,采集其花药作为实验材料,以前人研究得出的结果,SQ-1诱导的小麦雄性不育大多发生在单核后期至二核期,故以处理和对照花药的二核期cDNA材料为基础,通过NCBI查询拟南芥SKP1基因的序列从而设计引物并进行SKP1基因全长的扩增,将分别从SQ-1处理的和未处理的二核期材料中扩增得到的序列进行序列同源性比对,同时,进行西农2611花药处于单核期,二核期,三核期各时期SKP1基因的荧光定量表达,试图为揭示小麦生理型雄性不育的机理奠定一定理论基础。通过研究,获得如下结果:
(1)SKP1基因的表达与SQ-1诱导的小麦生理型雄性不育相关。
(2)从西农2611处理的和未处理的二核期cDNA中扩增,五次重复测序,最终得到了一段852bp长度的基因序列,经过比对和判断是属于小麦SKP1基因家族中的一份子,从而得到了新的小麦西农2611二核期SKP1基因序列,并将其上传至NCBI文库。
(3)分别所获得的SQ-1处理的二核期和对照二核期2611的两段全长基因序列利用软件DNAman进行序列比对,结果发现,这两段基因序列的比对结果显示,只有第477个碱基有差异,其余的碱基序列完全保持一致。将此二者基因序列利用PrimerPrimer5.0软件进行翻译表达并比较二者翻译的蛋白质序列,结果得出,一个碱基差异并不能引起蛋白质翻译的差异。由此看出,在经过SQ-1处理的材料和对照材料中,SKP1基因的转录并没有受到影响,也就是说,SQ-1处理小麦未能引起基因转录水平的差异。
(4)利用荧光定量PCR技术对SKP1基因在生理型不育株和可育株花药中表达进行了定量分析。结果表明,该基因在不育株和可育株花药发育3个时期(单核、二核和三核)表达量均表现为下降趋势,并且该基因在不育株花药发育的三个时期表达均相对于可育株被明显抑制。推测该基因在花粉粒发育过程中下调表达可能影响了小麦花粉粒发育过程中蛋白质泛素降解途径而导致雄性不育。
The whea(tTriticum aestivum)is very impotant grain in world, also one of main material basis to our existing.It’s an effective measure of increasing production and improving quality by heterosis. However, The theory of heterosis and research of practice with wheat are not deeper than maize, rice, rape and other crops.So far, study on the mechanism of male sterility is a significant breakthrough point for using heterosis.Summarized domestic and international wheat infertility type, It has two major aspects for studing male sterility including: male-sterility induced by Physiology and heredity. In the research, different scholars have different emphasis, they do these series study including the fertility of genes, difference of fertility proteins,physiological and biochemical metabolism. Ubiquitin proteasome pathway which can guarantee complex physilolgical and biochemical processes in proper operation is an important way to degradate redundancy protein in cells.One kind of low molecular weight protein SKP1(S-phase kinase-associated-protein 1)participating in ubiquitin proteasome pathway is found existing in eukaryotes broadly.It plays a decisive role in maintaining the biochemical function of SCF complex.The process of protein-degradating of ubiquitin mediated in cells is mainly regulated by SCF related with regulation in cell cycle.And at the same time, many physiological processes depends on SCF complex.
In the study, male sterility intruduced by chemical hybrid agent SQ-1, the anther of xinong-2611 divided into two samples-the spraying and unspraying as the test object. From the former study,we find that male sterile induced by SQ-1 happens from the later of mononuclear to binuclear stage.So take cDNA of binuclear stage of the two samples as the test object,and then the primer of SKP2 gene is designed and amplificated based on sequence of SKP1 gene of arabidopsis from NCBI GeneBank.Next, make a homology comparison of the above sequence,and at the same time ,make an expression of quantification of SKP1 gene during mononuclear stage,binuclear stage and trinuclear stage,respectively in 2611.The study will lay the theoretical foundation for revealing the mechanism of male-sterile induced by physiology. The result is as follows via the experiment:
(1)expression of SKP1 is related with physiological type male sterility by SQ-1 treated in wheat
(2)used total RNA extracted from anthers in binuclear to strand cDNA by revearse transcrition,and then gene sequences(852bp) are finally gained. by comparison and judgement,the SKP1 gene sequence in binuclear stage of wheat2611 is found firstly and would to been submitted to NCBI GeneBank
(3)Making a comparison of the two sequences achieved above with DNAman software,the result reveals that all bases remain unchanged except for the No.477 base.Through contrasting the protein sequences translated from the different bases with PrimerPrimer5.0 software,only one different base can’t cause difference of protein.Conclusion: transcription of SKP1 gene remains uninfluenced on both test object-the spraying and unspraying, in other words,SQ-1 can’t cause difference at transcriptional level on wheat.
(4)Aalysis the expression of SKP1 gene of anthers in sterile plant and fertile plant induced by physiology respectively via fluorescence quantitatiye PCR technique. The results suggest that the expression of the gene all appears an decreasing tendency in three periods(mononuclear, binuclear and trinuclear)during the development of anthers in sterile and fertile plant. It’s obvious that the expression of the gene in sterility plant during this three periods was dramatically inhibited compared with that in fertile plant. As a result,it can be easily supposed that influencing the ubiquitin-proteasome proteolytic pathway in the development of pollen grains,down-regulated SKP1 gene expression can lead to male sterility.
(1)SKP1基因的表达与SQ-1诱导的小麦生理型雄性不育相关。
(2)从西农2611处理的和未处理的二核期cDNA中扩增,五次重复测序,最终得到了一段852bp长度的基因序列,经过比对和判断是属于小麦SKP1基因家族中的一份子,从而得到了新的小麦西农2611二核期SKP1基因序列,并将其上传至NCBI文库。
(3)分别所获得的SQ-1处理的二核期和对照二核期2611的两段全长基因序列利用软件DNAman进行序列比对,结果发现,这两段基因序列的比对结果显示,只有第477个碱基有差异,其余的碱基序列完全保持一致。将此二者基因序列利用PrimerPrimer5.0软件进行翻译表达并比较二者翻译的蛋白质序列,结果得出,一个碱基差异并不能引起蛋白质翻译的差异。由此看出,在经过SQ-1处理的材料和对照材料中,SKP1基因的转录并没有受到影响,也就是说,SQ-1处理小麦未能引起基因转录水平的差异。
(4)利用荧光定量PCR技术对SKP1基因在生理型不育株和可育株花药中表达进行了定量分析。结果表明,该基因在不育株和可育株花药发育3个时期(单核、二核和三核)表达量均表现为下降趋势,并且该基因在不育株花药发育的三个时期表达均相对于可育株被明显抑制。推测该基因在花粉粒发育过程中下调表达可能影响了小麦花粉粒发育过程中蛋白质泛素降解途径而导致雄性不育。
The whea(tTriticum aestivum)is very impotant grain in world, also one of main material basis to our existing.It’s an effective measure of increasing production and improving quality by heterosis. However, The theory of heterosis and research of practice with wheat are not deeper than maize, rice, rape and other crops.So far, study on the mechanism of male sterility is a significant breakthrough point for using heterosis.Summarized domestic and international wheat infertility type, It has two major aspects for studing male sterility including: male-sterility induced by Physiology and heredity. In the research, different scholars have different emphasis, they do these series study including the fertility of genes, difference of fertility proteins,physiological and biochemical metabolism. Ubiquitin proteasome pathway which can guarantee complex physilolgical and biochemical processes in proper operation is an important way to degradate redundancy protein in cells.One kind of low molecular weight protein SKP1(S-phase kinase-associated-protein 1)participating in ubiquitin proteasome pathway is found existing in eukaryotes broadly.It plays a decisive role in maintaining the biochemical function of SCF complex.The process of protein-degradating of ubiquitin mediated in cells is mainly regulated by SCF related with regulation in cell cycle.And at the same time, many physiological processes depends on SCF complex.
In the study, male sterility intruduced by chemical hybrid agent SQ-1, the anther of xinong-2611 divided into two samples-the spraying and unspraying as the test object. From the former study,we find that male sterile induced by SQ-1 happens from the later of mononuclear to binuclear stage.So take cDNA of binuclear stage of the two samples as the test object,and then the primer of SKP2 gene is designed and amplificated based on sequence of SKP1 gene of arabidopsis from NCBI GeneBank.Next, make a homology comparison of the above sequence,and at the same time ,make an expression of quantification of SKP1 gene during mononuclear stage,binuclear stage and trinuclear stage,respectively in 2611.The study will lay the theoretical foundation for revealing the mechanism of male-sterile induced by physiology. The result is as follows via the experiment:
(1)expression of SKP1 is related with physiological type male sterility by SQ-1 treated in wheat
(2)used total RNA extracted from anthers in binuclear to strand cDNA by revearse transcrition,and then gene sequences(852bp) are finally gained. by comparison and judgement,the SKP1 gene sequence in binuclear stage of wheat2611 is found firstly and would to been submitted to NCBI GeneBank
(3)Making a comparison of the two sequences achieved above with DNAman software,the result reveals that all bases remain unchanged except for the No.477 base.Through contrasting the protein sequences translated from the different bases with PrimerPrimer5.0 software,only one different base can’t cause difference of protein.Conclusion: transcription of SKP1 gene remains uninfluenced on both test object-the spraying and unspraying, in other words,SQ-1 can’t cause difference at transcriptional level on wheat.
(4)Aalysis the expression of SKP1 gene of anthers in sterile plant and fertile plant induced by physiology respectively via fluorescence quantitatiye PCR technique. The results suggest that the expression of the gene all appears an decreasing tendency in three periods(mononuclear, binuclear and trinuclear)during the development of anthers in sterile and fertile plant. It’s obvious that the expression of the gene in sterility plant during this three periods was dramatically inhibited compared with that in fertile plant. As a result,it can be easily supposed that influencing the ubiquitin-proteasome proteolytic pathway in the development of pollen grains,down-regulated SKP1 gene expression can lead to male sterility.
引文
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