中国石蒜离体培养过程中腋芽发生的发育学机理研究
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摘要
石蒜属植物是一类集中分布在中国等东亚国家的多年生鳞茎类草本植物。独特的花型、多变的花色、丰富的加兰他敏和淀粉含量使之具有巨大的经济开发价值。本研究选取石蒜属中淀粉和药用价值均较高、全国广布的一个种—中国石蒜为研究材料，在建立起完整的离体培养体系后，从组织结构、激素、特异蛋白质和基因表达等多个层面深入探索腋芽发生的发育规律；并初步提出了改进离体培养的操作建议。主要结果如下：
1.以中国石蒜带基盘双鳞叶为外植体，经过诱导腋芽、增殖腋芽、生根等发育阶段后，形成完整的再生植株。外源细胞分裂素对启动腋芽发生、发育起到关键作用，并在一定范围内促进腋芽再生。在此基础上，对植株再生过程用石蜡切片和扫描电镜进行了组织学观察。结果表明本研究所建立的中国石蒜的离体培养过程属于定芽发生体系，表现出稳定的诱导率和增殖率。
2.在形态学观察的基础上，用两种分子标记-ISSR、SRAP-对连续培养27个月的再生植株进行了体细胞无性系变异研究。结果显示，两种分子标记均呈现出极低的变异率，说明建立起的离体培养体系可以较好的保存种质资源。
3.用POX、AMY和ATP三种同工酶从蛋白质水平对离体培养过程的发育机理进行了初步研究。结果表明，三种同工酶在不同发育阶段发挥不同的作用，其变化规律与形态发生过程相吻合。
4.以CKs、IAA、GAs和ABA四种内源激素为指标，集中探究了这四种激素在中国石蒜植株再生过程中的作用模式和机制。结果表明，4种激素在不同发育阶段发挥的作用各不相同，互有配合或拮抗。外源细胞分裂素作为刺激信号是启动腋芽发生、维持植株再生的关键因素；内源细胞分裂素通过改变组织的生理状态使其有利于腋芽再生。同时，激素之间的互作用可能影响下游器官发生基因的表达，这二者是离体形态建成的主要原因。
5.为了从基因水平研究植株再生的发育规律，本研究在参考多糖、多淀粉植物RNA提取方法的基础上建立了适合石蒜属植物的RNA提取方法。方法的有效性经过石蒜属5个种不同类型材料的验证。
6.应用同源克隆法（RACE）从中国石蒜鳞叶中克隆出KNOX基因全长cDNA。序列分析表明，该基因在氨基酸水平上与多种单子叶植物的KNOX基因有较高的同源性。
7.应用荧光定量PCR对从中国石蒜鳞叶中克隆出LcKNOX1在离体培养过程中11个典型发育时间点的表达量进行了初步研究，结果表明内源CKs的含量与该基因的表达量有极显著正相关性，并做出了二者之间的直线回归方程。综合内源激素和基因表达数据，我们推测外源细胞分裂素作为刺激信号诱使内源细胞分裂素积累并与其它激素一起促使形态建成。最后，结合激素和基因表达结果，我们提出了改进离体培养的初步建议。
Lycoris, mainly distribute in China, is a peculiar genus in monocots. It is diversity in flowerpattern as well as colors and also abundant in galanthamin and starch, which made Lycoris itselfgreat economic value.
In view of morphogenesis, endogenous hormones, isozymes and gene, the developmentprocess of axillary bud regeneration was specifically studied in this research. And the mainresults are as follows:
1. A complete definite-bud regeneration in vitro culture system with stable induction rateand proliferation rate was established through the induction, proliferation and rooting mediums.Extrogenous-cytokinin played an important role in the initiation of axillary bud and itssubsequent development. The morphogenesis process was described by serial sections andscanning electron microscope (SEM) in details.
2. Genetic variability of micropropagated plantlets, also called somaclonal variation, wasassessed by two different PCR-based markers, ISSRs (inter-simple sequence repeats) and SRAPs(sequence-related amplified polymorphisms) markers. Both markers showed extremely lowmutation rates, which suggested that the established system is appropriate for clonal propagationand germplasm of Lycoris chinensis preservation.
3. The mechanism of in vitro regeneration was sketchily discussed by3isozymes atprotein level in this paper. The results showed that there is a correlation between structuralchanges and protein activity and was proved by POX, AMY and ATP isozymes.
4. CKs、IAA、GAs and ABA were token as judgement indexes to analyze the function ofendogenous-hormones. The results showed that these endo-hormones played different roles in thecourse of organogenesis development. Endo-cytokinin and auxin interacted together to changethe physiological status. They promoted “axillary bud primordium” transiting from “dormancystate” to “development state”, organ initiation and morphogenesis. The interactions among theseendogenous-hormones affected the expression of downstream organogenesis genes and proteins.
5. A proper RNA extraction method was established for analyzing the axillary buddevelopment mechanism at gene level. Efficiency of this method was testified by3kinds ofmaterials of5species of Lycoris.
6. Complete-length cDNAs encoding KNOX was isolated from the bulb of L.chinensis byRACE. Bioinformatics analysis showed that the cloned LcKNOX1shared a high similarity withperviously reported monocotyledon KNOX protein.
7. The expression profiles of LcKNOX1at11sampling times were analyzed byquantitative real time PCR. The results showed that there was a significant linear positivecorrelation between the level of CKs and the expression of LcKNOX1. That is to say, theconcentration of endo-cytokinin was directly associated with the elevation of expression ofLcKNOX1. Meanwhile, based on the results of qPCR and endogenous hormones, preliminary suggestions of reverse guidance for improving the in vitro culture process was proposed.
1.以中国石蒜带基盘双鳞叶为外植体，经过诱导腋芽、增殖腋芽、生根等发育阶段后，形成完整的再生植株。外源细胞分裂素对启动腋芽发生、发育起到关键作用，并在一定范围内促进腋芽再生。在此基础上，对植株再生过程用石蜡切片和扫描电镜进行了组织学观察。结果表明本研究所建立的中国石蒜的离体培养过程属于定芽发生体系，表现出稳定的诱导率和增殖率。
2.在形态学观察的基础上，用两种分子标记-ISSR、SRAP-对连续培养27个月的再生植株进行了体细胞无性系变异研究。结果显示，两种分子标记均呈现出极低的变异率，说明建立起的离体培养体系可以较好的保存种质资源。
3.用POX、AMY和ATP三种同工酶从蛋白质水平对离体培养过程的发育机理进行了初步研究。结果表明，三种同工酶在不同发育阶段发挥不同的作用，其变化规律与形态发生过程相吻合。
4.以CKs、IAA、GAs和ABA四种内源激素为指标，集中探究了这四种激素在中国石蒜植株再生过程中的作用模式和机制。结果表明，4种激素在不同发育阶段发挥的作用各不相同，互有配合或拮抗。外源细胞分裂素作为刺激信号是启动腋芽发生、维持植株再生的关键因素；内源细胞分裂素通过改变组织的生理状态使其有利于腋芽再生。同时，激素之间的互作用可能影响下游器官发生基因的表达，这二者是离体形态建成的主要原因。
5.为了从基因水平研究植株再生的发育规律，本研究在参考多糖、多淀粉植物RNA提取方法的基础上建立了适合石蒜属植物的RNA提取方法。方法的有效性经过石蒜属5个种不同类型材料的验证。
6.应用同源克隆法（RACE）从中国石蒜鳞叶中克隆出KNOX基因全长cDNA。序列分析表明，该基因在氨基酸水平上与多种单子叶植物的KNOX基因有较高的同源性。
7.应用荧光定量PCR对从中国石蒜鳞叶中克隆出LcKNOX1在离体培养过程中11个典型发育时间点的表达量进行了初步研究，结果表明内源CKs的含量与该基因的表达量有极显著正相关性，并做出了二者之间的直线回归方程。综合内源激素和基因表达数据，我们推测外源细胞分裂素作为刺激信号诱使内源细胞分裂素积累并与其它激素一起促使形态建成。最后，结合激素和基因表达结果，我们提出了改进离体培养的初步建议。
Lycoris, mainly distribute in China, is a peculiar genus in monocots. It is diversity in flowerpattern as well as colors and also abundant in galanthamin and starch, which made Lycoris itselfgreat economic value.
In view of morphogenesis, endogenous hormones, isozymes and gene, the developmentprocess of axillary bud regeneration was specifically studied in this research. And the mainresults are as follows:
1. A complete definite-bud regeneration in vitro culture system with stable induction rateand proliferation rate was established through the induction, proliferation and rooting mediums.Extrogenous-cytokinin played an important role in the initiation of axillary bud and itssubsequent development. The morphogenesis process was described by serial sections andscanning electron microscope (SEM) in details.
2. Genetic variability of micropropagated plantlets, also called somaclonal variation, wasassessed by two different PCR-based markers, ISSRs (inter-simple sequence repeats) and SRAPs(sequence-related amplified polymorphisms) markers. Both markers showed extremely lowmutation rates, which suggested that the established system is appropriate for clonal propagationand germplasm of Lycoris chinensis preservation.
3. The mechanism of in vitro regeneration was sketchily discussed by3isozymes atprotein level in this paper. The results showed that there is a correlation between structuralchanges and protein activity and was proved by POX, AMY and ATP isozymes.
4. CKs、IAA、GAs and ABA were token as judgement indexes to analyze the function ofendogenous-hormones. The results showed that these endo-hormones played different roles in thecourse of organogenesis development. Endo-cytokinin and auxin interacted together to changethe physiological status. They promoted “axillary bud primordium” transiting from “dormancystate” to “development state”, organ initiation and morphogenesis. The interactions among theseendogenous-hormones affected the expression of downstream organogenesis genes and proteins.
5. A proper RNA extraction method was established for analyzing the axillary buddevelopment mechanism at gene level. Efficiency of this method was testified by3kinds ofmaterials of5species of Lycoris.
6. Complete-length cDNAs encoding KNOX was isolated from the bulb of L.chinensis byRACE. Bioinformatics analysis showed that the cloned LcKNOX1shared a high similarity withperviously reported monocotyledon KNOX protein.
7. The expression profiles of LcKNOX1at11sampling times were analyzed byquantitative real time PCR. The results showed that there was a significant linear positivecorrelation between the level of CKs and the expression of LcKNOX1. That is to say, theconcentration of endo-cytokinin was directly associated with the elevation of expression ofLcKNOX1. Meanwhile, based on the results of qPCR and endogenous hormones, preliminary suggestions of reverse guidance for improving the in vitro culture process was proposed.
引文
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