谷精草属植物个体早期发育及种间亲缘关系的研究
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摘要
谷精草属(Eriocaulon Linn.)植物属于湿生单子叶植物,其特殊的湿生生存条件及植株小的特点为其个体的早期发育研究造成了障碍,迄今,其种子萌发和幼苗发育的研究资料甚少,发育过程中的解剖结构描述不清楚,存在较大争议。同时,由于种内个体变异较大,种间相似度较高,导致分类和系统研究的困难,至今仍没有形成统一认识。针对上述问题,本文通过对谷精草属种子萌发过程中的生理和解剖学研究,探讨其个体早期发育的特点;将分子生物学技术和形态学研究相结合,对谷精草属的6个组10个种进行了同工酶分子标记和扩增片段长度多态性分子标记(AFLP)的分析,探讨其系统发育关系及种间遗传多样性,丰富了该属植物个体发育及系统进化研究的数据及理论。主要结果如下:
1.种子萌发研究结果表明,谷精草种子具有需高温、高湿和非光敏性的萌发习性。
2.采用光学、透射电子、扫描电子、激光共聚焦及荧光显微镜等对种子萌发过程中形态结构的变化情况进行研究。结果表明,谷精草作为湿生植物,为适应环境,器官的发生在顺序上与陆生植物不同,并出现一些特殊结构,具有以下特点:⑴存在胚的后发育过程,即随着种子萌发,胚的各部分逐渐分化完全;⑵没有明显的下胚轴,根区组织突破种皮主要靠根区细胞伸长作用及顶细胞分生组织的不断分化;⑶幼苗上着生多种表皮毛,根区表皮细胞发育成单细胞毛,胚芽鞘基部着生多细胞表皮毛。
3.对谷精草属6个种的过氧化物酶(POD)、酯酶(EST)、苹果酸脱氢酶(MDH)、淀粉酶(AMY)的同工酶进行了分离,采用STATISTICA/W 4.5软件对酶谱的不同相似性系数进行聚类分析,绘出树状图。结果显示E. alpestre同E. buergerianum聚为一枝,与E. sexangulare聚为一组,而E. nepalense、E. kunmingense、E. cinereum聚为一组,E. kunmingense同E. cinereum关系较近。
4.以谷精草属10个种进行AFLP标记,利用8对选择性扩增引物组合,共扩增产生1555条谱带,其中1552条谱带表现出多态性(99.81%),说明谷精草属植物种间具有丰富的遗传多样性。用NTSYSpc-2.11F计算所得的[0,1]原始矩阵的J & C距离。将所得遗传距离矩阵转换为meg文件格式,在MEGA(Molecular EvolutionaryGenetics Analysis)中以最小进化(Minimum Evolution)法生成系统发育树。结果显示E. alpestre同E. buergerianum;E. cinereum同E. kunmingense聚在一起。
同工酶酶谱及AFLP标记聚类分析的结果,与形态学研究相比较,支持将昆明谷精草(E. kunmingense Z. X. Zhang)同白药谷精草(E. cinereum R. Brown.)归为一组,与谷精草属近期报道的分类系统相符,为进一步确定谷精草属和谷精草科的系统进化位置提供了分子水平的实验依据。
Eriocaulon Linn., belonging to hygrophyte monocotyledons, is difficulty to research on the stage of ontogenesis, because of their aquatic living conditions and tiny figures. Up to now, the studies on the seed germinates and ontogenesis remain limited. Therefore, the anatomic structures in the process of germinates are still obscure. At the same time, the high intraspecific variability combined within the limited interspecific differences makes the taxonomy of Eriocaulon Linn. very difficult. Therefore, there are many debates on this field. In this paper, the physiological and anatomic researches on seeds germination were carried out to study the early stage of ontogeny of Eriocaulon. In addition, isozyme and AFLP molecular biotechnology were made a use of the analysis on the interspecific and phylogenetic relationships of 10 species of Eriocaulon. The results were shown as following:
1. The result indicated that the suitable condition of the seeds germination of Eriocaulon was high temperature, high humidity with non-photosensitive.
2. The result of study on the morphology of ontogenesis by using the light microscopy, TEM, SEM, LSCM and fluorescence-microscope indicated that, the Eriocaulon, as an aquatic, their organs with some particular structures to adapt to the environment were quite different from terrestrial plants. In conclusion:⑴There was no obvious differentiation in embryos, and both the shoot apex and the root apex developed very weakly after germination;⑵There was no classical hypocotyl, so radicle zone from the basal cell push out the micropyle by elongating themselves and the dividing of the organs from apical cell;⑶It was observed that the seedling had two kinds of epidermal hairs: the surface cells of radicular end developed to single-cellular hairs, while in the basal of the embryo sheaths existed multi-cellular hairs.
3. Four isozymes (POD、EST、MDH、AMY) in seeds of Eriocaulon Linn. were researched. Software STATISTICA/w 4.5 was used to analyze the result of electrophoresis. After that, a tree diagram of these 6 species was abained which showed a closer relationship between E. alpestre, E. buergerianum and E. sexangulare, while E. cinereum, E. nepalense and E. kunmingense to be another branch.
4. The results of AFLP research on 10 species showed that 1552 bands (99.81 %) behaved polymorphic in 1555 AFLP bands amplified by 8 primers-pairs. Using NTSYSpc-2.11F computer programs, The J & C distances of the data of 1555 AFLP bands were generated and a dendrogram was constructed, which showed that there was a nearer phylogenetic relationship between E. alpestre and E. buergerianum, as while as E. cinereum and E. kunmingense.
The results of cluster analysis about isozyme and AFLP, comparing with morphology study, supports the view that E. kunmingense Z. X. Zhang and E. cinereum R. Brown should be put into one section, which confirmed to the latest taxonomy system of Eriocaulon Linn.. Further more, it would provide basis for the further study on the confirmation of the phylogenetic position of Eriocaulaceae and Eriocaulon.
1.种子萌发研究结果表明,谷精草种子具有需高温、高湿和非光敏性的萌发习性。
2.采用光学、透射电子、扫描电子、激光共聚焦及荧光显微镜等对种子萌发过程中形态结构的变化情况进行研究。结果表明,谷精草作为湿生植物,为适应环境,器官的发生在顺序上与陆生植物不同,并出现一些特殊结构,具有以下特点:⑴存在胚的后发育过程,即随着种子萌发,胚的各部分逐渐分化完全;⑵没有明显的下胚轴,根区组织突破种皮主要靠根区细胞伸长作用及顶细胞分生组织的不断分化;⑶幼苗上着生多种表皮毛,根区表皮细胞发育成单细胞毛,胚芽鞘基部着生多细胞表皮毛。
3.对谷精草属6个种的过氧化物酶(POD)、酯酶(EST)、苹果酸脱氢酶(MDH)、淀粉酶(AMY)的同工酶进行了分离,采用STATISTICA/W 4.5软件对酶谱的不同相似性系数进行聚类分析,绘出树状图。结果显示E. alpestre同E. buergerianum聚为一枝,与E. sexangulare聚为一组,而E. nepalense、E. kunmingense、E. cinereum聚为一组,E. kunmingense同E. cinereum关系较近。
4.以谷精草属10个种进行AFLP标记,利用8对选择性扩增引物组合,共扩增产生1555条谱带,其中1552条谱带表现出多态性(99.81%),说明谷精草属植物种间具有丰富的遗传多样性。用NTSYSpc-2.11F计算所得的[0,1]原始矩阵的J & C距离。将所得遗传距离矩阵转换为meg文件格式,在MEGA(Molecular EvolutionaryGenetics Analysis)中以最小进化(Minimum Evolution)法生成系统发育树。结果显示E. alpestre同E. buergerianum;E. cinereum同E. kunmingense聚在一起。
同工酶酶谱及AFLP标记聚类分析的结果,与形态学研究相比较,支持将昆明谷精草(E. kunmingense Z. X. Zhang)同白药谷精草(E. cinereum R. Brown.)归为一组,与谷精草属近期报道的分类系统相符,为进一步确定谷精草属和谷精草科的系统进化位置提供了分子水平的实验依据。
Eriocaulon Linn., belonging to hygrophyte monocotyledons, is difficulty to research on the stage of ontogenesis, because of their aquatic living conditions and tiny figures. Up to now, the studies on the seed germinates and ontogenesis remain limited. Therefore, the anatomic structures in the process of germinates are still obscure. At the same time, the high intraspecific variability combined within the limited interspecific differences makes the taxonomy of Eriocaulon Linn. very difficult. Therefore, there are many debates on this field. In this paper, the physiological and anatomic researches on seeds germination were carried out to study the early stage of ontogeny of Eriocaulon. In addition, isozyme and AFLP molecular biotechnology were made a use of the analysis on the interspecific and phylogenetic relationships of 10 species of Eriocaulon. The results were shown as following:
1. The result indicated that the suitable condition of the seeds germination of Eriocaulon was high temperature, high humidity with non-photosensitive.
2. The result of study on the morphology of ontogenesis by using the light microscopy, TEM, SEM, LSCM and fluorescence-microscope indicated that, the Eriocaulon, as an aquatic, their organs with some particular structures to adapt to the environment were quite different from terrestrial plants. In conclusion:⑴There was no obvious differentiation in embryos, and both the shoot apex and the root apex developed very weakly after germination;⑵There was no classical hypocotyl, so radicle zone from the basal cell push out the micropyle by elongating themselves and the dividing of the organs from apical cell;⑶It was observed that the seedling had two kinds of epidermal hairs: the surface cells of radicular end developed to single-cellular hairs, while in the basal of the embryo sheaths existed multi-cellular hairs.
3. Four isozymes (POD、EST、MDH、AMY) in seeds of Eriocaulon Linn. were researched. Software STATISTICA/w 4.5 was used to analyze the result of electrophoresis. After that, a tree diagram of these 6 species was abained which showed a closer relationship between E. alpestre, E. buergerianum and E. sexangulare, while E. cinereum, E. nepalense and E. kunmingense to be another branch.
4. The results of AFLP research on 10 species showed that 1552 bands (99.81 %) behaved polymorphic in 1555 AFLP bands amplified by 8 primers-pairs. Using NTSYSpc-2.11F computer programs, The J & C distances of the data of 1555 AFLP bands were generated and a dendrogram was constructed, which showed that there was a nearer phylogenetic relationship between E. alpestre and E. buergerianum, as while as E. cinereum and E. kunmingense.
The results of cluster analysis about isozyme and AFLP, comparing with morphology study, supports the view that E. kunmingense Z. X. Zhang and E. cinereum R. Brown should be put into one section, which confirmed to the latest taxonomy system of Eriocaulon Linn.. Further more, it would provide basis for the further study on the confirmation of the phylogenetic position of Eriocaulaceae and Eriocaulon.
引文
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