苜蓿花药培养与雄性不育差异表达分析
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摘要
为了解雄性不育形成的分子机理,获得苜蓿雄性不育纯合系植株,进一步提高苜蓿雄性不育杂种优势的利用价值,本试验以苜蓿雄性不育株及其对应可育株的花蕾作为研究对象,通过花药组织培养技术获得苜蓿单倍体植株,同时利用cDNA.AFLP差异显示技术对不同发育时期不育株和可育株花蕾的基因差异表达进行研究,探讨了苜蓿雄性不育性的基因差异表达模式,并进一步对差异基因片段进行氨基酸序列和蛋白质结构的分析。主要研究结果如下：
(1)苜蓿雄性不育株与可育株花药组织培养的研究表明,在现蕾初期取材、不经低温处理的花药愈伤组织诱导率可高达66.7%,且培养条件为MS+2,4-D0.5mg/L+6-BA0.25mg/L+NAA0.2mg/L+KT3mg/L+3%蔗糖+0.7%琼脂：适宜不育材料的分化培养基为MS+KT1mg/L+NAA0.2mg/L+2%蔗糖+0.7%琼脂,适宜可育材料的为MS+KT4mg/L+NAA0.2mg/L+2%蔗糖+0.7%琼脂；适合诱导幼苗生根的培养基为1/2MS+NAA0.1mg/L+2%蔗糖+0.9%琼脂；此外,不添加激素的MS培养基可以诱导玻璃化苗生根。
(2)在苜蓿花药组织培养再生植株的倍性鉴定研究中,适宜的细胞核悬液制备方法是选择新鲜幼苗叶片,在0.1mol/L柠檬酸和0.2%TritonX.100组成的分离缓冲液中用刀片进行机械切割,经两步离心法(800rpm离心5min)分离；同时,利用100～200μL浓度为50mg/L的PI染液对材料细胞核染色效果好；通过流式细胞仪对再生植株进行倍性鉴定,结果获得单倍体、四倍体和混倍体的比例分别为：16%、78%和6%。
(3)对苜蓿不育株与可育株花蕾的cDNA.AFLP技术体系进行优化,发现该技术重复性高、多态性好,适用于苜蓿不同时期花蕾基因差异表达的研究。通过12对引物获得1746条能够稳定存在的条带,其中差异表达基因占12.37%,且包括不育性特异表达、不育性表达和育性特异表达3种类型。
(4)将4条苜蓿花蕾期的基因差异片段进行cDNA序列分析。通过NCBI中BlastN和BlastX数据库检索,结果发现3条片段与苜蓿具有较高同源性(93%～100%)。其中,3号差异片段与蚕豆叶绿体中的1,5-二磷酸核酮糖羧化基因和氧化酶大亚基基因的同源性高达98～99%,而二磷酸核酮塘羧化酶与植物的CMS有关,参与了小麦花药发育调控、细胞程序性死亡及物质能量代谢等过程,因此推测该差异片段与苜蓿雄性不育性的形成有关。
Research on the formation mechanism of male sterility in alfalfa is important for obtaining the male sterility pure lines and improving the heterosis utilization of alfalfa. In this study, the bud of male sterile and fertile plants of alfalfa was considered as the experimental materils. The anther culture technique was used to cultivate alfalfa haploid plants. And the cDNA-AFLP differential display technique were used to research on the level of gene regulation in different growth period of alfalfa bud, and to discuss its gene differential expression patterns of alfalfa male sterility and analyse the amino acid sequence and protein structure of different genes, in order to analyse the formation mechanism of male sterility. The main results were as follows:
(1) The study on male sterile and fertile line of alfalfa anther culture showed that the good callus induction can get under the condition of sampling materials in the bud stage, without per-treatment with low temperature, and the highest callus induction rate was66.7%. MS+2,4-D0.5mg/L+6-BA0.25mg/L+NAA0.2mg/L+KT3mg/L+sucrose3%+ager0.7%was suited for callus induction. MS+KT lmg/L+NAA0.2mg/L+sucrose2%+ager0.7%and MS+KT4mg/L+NAA0.2mg/L+sucrose2%+ager0.7%was suited for sterile and fertile callus differentiation respectively.1/2MS+NAA0.1mg/L+sucrose2%+ager0.9%was suited for rooting. Furthermore, the vitreous shoots could be rooted by MS.
(2) DNA contents and ploidy of alfalfa anther culture plants were detected by flow-cytometric analysis. The proper cell nucleus suspensions could be obtained by cutting the fresh leaves in the suitable buffer solution (0.1mol/L citric acid and0.2%TritonX-100) and centrifuging twice (800rpm,5min). Moreover, the most appropriate dyeing effects to cell nucleus were realized by PI (100～200μL,50mg/L). In a word, the results of flow-cytometric indicated that16%plants were haploid,78%plants were tetraploid, and6%plants were mixoploid.
(3) The buds gene differential expression of afalfa male sterile and fertile lines were analyzed by cDNA-AFLP technology. The results showed that cDNA-AFLP could be applicable to the research on differential gene expression of different alfalfa buds stages. In addition, its had a high reproducibility and good polymorphism.1746stable bands were detected with12primers, and proportion of differentially expressed genes was12.37%which including3differential gene expression patterns (sterility specific expression, sterility expression, and fertility specific expression).
(4) Through identification of repeated tests,4alfalfa buds gene fragments were randomly cloned and analysed by BlastN and BlastX from NCBI. The retrieval results were indicated that3gene fragments had a high homologous with alfalfa (93%～100%). Especially, No.3differential fragment had98%～99%nucleotide similarity to1,5-bisphosphate carboxylase/oxygenase large subunit gene of broad bean chloroplast. What's more, the CMS was controlled by the1,5-bisphosphate carboxylase, which involved in anther development, programmed cell death, and material energy metabolism of wheat. After sequence analysis, it found that the No.3differential fragment was maybe related to male sterility in alfalfa.
(1)苜蓿雄性不育株与可育株花药组织培养的研究表明,在现蕾初期取材、不经低温处理的花药愈伤组织诱导率可高达66.7%,且培养条件为MS+2,4-D0.5mg/L+6-BA0.25mg/L+NAA0.2mg/L+KT3mg/L+3%蔗糖+0.7%琼脂：适宜不育材料的分化培养基为MS+KT1mg/L+NAA0.2mg/L+2%蔗糖+0.7%琼脂,适宜可育材料的为MS+KT4mg/L+NAA0.2mg/L+2%蔗糖+0.7%琼脂；适合诱导幼苗生根的培养基为1/2MS+NAA0.1mg/L+2%蔗糖+0.9%琼脂；此外,不添加激素的MS培养基可以诱导玻璃化苗生根。
(2)在苜蓿花药组织培养再生植株的倍性鉴定研究中,适宜的细胞核悬液制备方法是选择新鲜幼苗叶片,在0.1mol/L柠檬酸和0.2%TritonX.100组成的分离缓冲液中用刀片进行机械切割,经两步离心法(800rpm离心5min)分离；同时,利用100～200μL浓度为50mg/L的PI染液对材料细胞核染色效果好；通过流式细胞仪对再生植株进行倍性鉴定,结果获得单倍体、四倍体和混倍体的比例分别为：16%、78%和6%。
(3)对苜蓿不育株与可育株花蕾的cDNA.AFLP技术体系进行优化,发现该技术重复性高、多态性好,适用于苜蓿不同时期花蕾基因差异表达的研究。通过12对引物获得1746条能够稳定存在的条带,其中差异表达基因占12.37%,且包括不育性特异表达、不育性表达和育性特异表达3种类型。
(4)将4条苜蓿花蕾期的基因差异片段进行cDNA序列分析。通过NCBI中BlastN和BlastX数据库检索,结果发现3条片段与苜蓿具有较高同源性(93%～100%)。其中,3号差异片段与蚕豆叶绿体中的1,5-二磷酸核酮糖羧化基因和氧化酶大亚基基因的同源性高达98～99%,而二磷酸核酮塘羧化酶与植物的CMS有关,参与了小麦花药发育调控、细胞程序性死亡及物质能量代谢等过程,因此推测该差异片段与苜蓿雄性不育性的形成有关。
Research on the formation mechanism of male sterility in alfalfa is important for obtaining the male sterility pure lines and improving the heterosis utilization of alfalfa. In this study, the bud of male sterile and fertile plants of alfalfa was considered as the experimental materils. The anther culture technique was used to cultivate alfalfa haploid plants. And the cDNA-AFLP differential display technique were used to research on the level of gene regulation in different growth period of alfalfa bud, and to discuss its gene differential expression patterns of alfalfa male sterility and analyse the amino acid sequence and protein structure of different genes, in order to analyse the formation mechanism of male sterility. The main results were as follows:
(1) The study on male sterile and fertile line of alfalfa anther culture showed that the good callus induction can get under the condition of sampling materials in the bud stage, without per-treatment with low temperature, and the highest callus induction rate was66.7%. MS+2,4-D0.5mg/L+6-BA0.25mg/L+NAA0.2mg/L+KT3mg/L+sucrose3%+ager0.7%was suited for callus induction. MS+KT lmg/L+NAA0.2mg/L+sucrose2%+ager0.7%and MS+KT4mg/L+NAA0.2mg/L+sucrose2%+ager0.7%was suited for sterile and fertile callus differentiation respectively.1/2MS+NAA0.1mg/L+sucrose2%+ager0.9%was suited for rooting. Furthermore, the vitreous shoots could be rooted by MS.
(2) DNA contents and ploidy of alfalfa anther culture plants were detected by flow-cytometric analysis. The proper cell nucleus suspensions could be obtained by cutting the fresh leaves in the suitable buffer solution (0.1mol/L citric acid and0.2%TritonX-100) and centrifuging twice (800rpm,5min). Moreover, the most appropriate dyeing effects to cell nucleus were realized by PI (100～200μL,50mg/L). In a word, the results of flow-cytometric indicated that16%plants were haploid,78%plants were tetraploid, and6%plants were mixoploid.
(3) The buds gene differential expression of afalfa male sterile and fertile lines were analyzed by cDNA-AFLP technology. The results showed that cDNA-AFLP could be applicable to the research on differential gene expression of different alfalfa buds stages. In addition, its had a high reproducibility and good polymorphism.1746stable bands were detected with12primers, and proportion of differentially expressed genes was12.37%which including3differential gene expression patterns (sterility specific expression, sterility expression, and fertility specific expression).
(4) Through identification of repeated tests,4alfalfa buds gene fragments were randomly cloned and analysed by BlastN and BlastX from NCBI. The retrieval results were indicated that3gene fragments had a high homologous with alfalfa (93%～100%). Especially, No.3differential fragment had98%～99%nucleotide similarity to1,5-bisphosphate carboxylase/oxygenase large subunit gene of broad bean chloroplast. What's more, the CMS was controlled by the1,5-bisphosphate carboxylase, which involved in anther development, programmed cell death, and material energy metabolism of wheat. After sequence analysis, it found that the No.3differential fragment was maybe related to male sterility in alfalfa.
引文
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