五味子地下横走茎发生机理研究
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摘要
本试验以五味子1年生、2年生和6年生植株为试材,研究了五味子地下横走茎、根、及地上茎形态解剖结构和不同节位腋芽亚显微结构的差异,并对其地下横走茎的形态发生进行了系统观察。通过对五味子生长过程中地上茎尖、地下横走茎茎尖等器官的可溶性糖、可溶性蛋白、内源激素含量的测定和过氧化物酶同工酶酶谱的分析,揭示了五味子地下横走茎发生发育与各因素间的内在联系。结果表明:
1.五味子地下横走茎与地上茎在发育过程中存在相似之处,都有原生分生组织、初生生长和次生生长等阶段。但初生生长在亚显微结构上有很大差别,地上茎尖第一层细胞外有明显薄膜、核膜发育完整、质体多、液泡相对较少,地下横走茎茎尖液泡多、营养多以脂滴形式存在;在茎尖形态、次生结构上也有明显差别,地下茎尖呈弯钩状,次生维管组织在横切面上的比例较地上茎小。
2.五味子根次生结构包括周皮、次生韧皮部、形成层和次生木质部,在次生生长的过程中,根中央有未木质化的薄壁细胞,地下横走茎初生木质部发育方式属于内始式,从本质上不同于根的初生结构发育方式。
3. 1年生五味子子叶腋芽可发育成地下横走茎,但其发育较缓慢,在8月下旬～9月初才发育成地下横走茎,植株的第1片真叶腋芽至第8片真叶腋芽在条件适合的情况下也可以发育成地下横走茎;地下横走茎的发生部位较集中,常呈轮生状;较高节位腋芽与子叶腋芽的亚显微结构存在较大差异,说明五味子的芽能否形成地下横走茎在其形成的早期可能就已经决定。
4.五味子地下横走茎在生长初期和休眠期都有较高的可溶性糖含量,这说明地下横走茎不仅有繁殖功能,而且有一定的贮藏功能,它与地上存在着营养竞争的关系;可溶性蛋白与生长的旺盛程度有关,在地下横走茎旺盛生长期,可溶性蛋白的含量也会相应地较高。
5.五味子地下横走茎发生发育过程中过氧化物酶(POD)在不同发育时期及部位存在较大差异。能发育成地下横走茎的芽比不能发育成地下横走茎的芽多两条酶带,可以将这两条带做为产生地下横走茎的特征酶带。
6.五味子植株的地下横走茎在发生初期ABA和ZR含量较高,说明其有利于地下横走茎的诱导,在伸长生长期则表现为IAA和GA3含量较高,说明这两种激素有利于地下横走茎的伸长生长。
In order to study the differences of morphology and anatomy between the root, stem and rhizome, one year old, two years old and six years old Schisandra chinensis (Turcz.) Baill were used as experiment materials and we also observed the morphogenesis of rhizome from seedling of Schisandra chinensis (Turcz.) Baill. Through the determination of soluble sugar, soluble protein, hormone and peroxidase isoenzyme in the process of formation of rhizome, we try to reveal the relations between the formation mechanism and these factors.The experiment results were as follows:
1. Development of the rhizome was similar to stem, both of them can be divided into promeristem, primary and secondary growth three stages. However, the ultrastructure of primary growth was different, for example, there was a special membrane clinging on the first layer cell of stem tip, and it was much more developed than rhizome tip with completed envelope, more plastids and less vacuoles, while the rhizome tip had more vacuoles and lipid droplets. Furthermore, they had other differences in tip type and secondary structure, for example, the rhizome tip was bendy and the proportion of secondary growth of the rhizome was less than stem.
2. Secondary structure of root was composed of periderm, cambium layer, secondary xylem and phloem, while we didn’t find the parenchyma cells with lignification, developing patterns of primary structure of rhizome is endarch, which is different of developing patterns of primary structure of root.
3. One year axillary bud of cotyledon could develop to rhizome in late August-early September slowly, and the axillary bud of rough leaf to the eighth euphylla would develop into rhizome finally when the condition was appropriate. Rotiform rhizomes were concentrated and whorled in the base of stem. Ultrastructure of axillary bud on the higher node have more differences from rough leaf, it would be decided in early stage that whether axillary bud of cotyledon develop to rhizome or not.
4. Rhizome tip of Schisandra chinensis (Turcz.) Baill had higher content of soluble sugar at initial and dormant period, which reflects the rhizome is different with other plants’rhizome. We found the rhizome not only had the reproductive function, but also had certain of storage feature, so the rhizome competed with the ground on nutrition. There was relationship between the content of soluble protein and the strong level of growth, and in the vigorous growth of rhizome, soluble protein content was consequently higher.
5. There was a great difference of peroxidase isoenzyme map in different developmental period and different type of buds in the process of the formation and development rhizome. The buds which can develop into rhizome have two more POD isoenzyme bands than the buds can’t develop into rhizome. The two POD isoenzyme bands is specific zymogram.
6. Levels of ABA and ZR were higher in formation period of rhizomes of Schisandra chinensis (Turcz.) Baill, which were conducive to induce rhizome. Levels IAA and GA3 were higher in the elongation growth stage, which were essential to rhizome elongation.
1.五味子地下横走茎与地上茎在发育过程中存在相似之处,都有原生分生组织、初生生长和次生生长等阶段。但初生生长在亚显微结构上有很大差别,地上茎尖第一层细胞外有明显薄膜、核膜发育完整、质体多、液泡相对较少,地下横走茎茎尖液泡多、营养多以脂滴形式存在;在茎尖形态、次生结构上也有明显差别,地下茎尖呈弯钩状,次生维管组织在横切面上的比例较地上茎小。
2.五味子根次生结构包括周皮、次生韧皮部、形成层和次生木质部,在次生生长的过程中,根中央有未木质化的薄壁细胞,地下横走茎初生木质部发育方式属于内始式,从本质上不同于根的初生结构发育方式。
3. 1年生五味子子叶腋芽可发育成地下横走茎,但其发育较缓慢,在8月下旬～9月初才发育成地下横走茎,植株的第1片真叶腋芽至第8片真叶腋芽在条件适合的情况下也可以发育成地下横走茎;地下横走茎的发生部位较集中,常呈轮生状;较高节位腋芽与子叶腋芽的亚显微结构存在较大差异,说明五味子的芽能否形成地下横走茎在其形成的早期可能就已经决定。
4.五味子地下横走茎在生长初期和休眠期都有较高的可溶性糖含量,这说明地下横走茎不仅有繁殖功能,而且有一定的贮藏功能,它与地上存在着营养竞争的关系;可溶性蛋白与生长的旺盛程度有关,在地下横走茎旺盛生长期,可溶性蛋白的含量也会相应地较高。
5.五味子地下横走茎发生发育过程中过氧化物酶(POD)在不同发育时期及部位存在较大差异。能发育成地下横走茎的芽比不能发育成地下横走茎的芽多两条酶带,可以将这两条带做为产生地下横走茎的特征酶带。
6.五味子植株的地下横走茎在发生初期ABA和ZR含量较高,说明其有利于地下横走茎的诱导,在伸长生长期则表现为IAA和GA3含量较高,说明这两种激素有利于地下横走茎的伸长生长。
In order to study the differences of morphology and anatomy between the root, stem and rhizome, one year old, two years old and six years old Schisandra chinensis (Turcz.) Baill were used as experiment materials and we also observed the morphogenesis of rhizome from seedling of Schisandra chinensis (Turcz.) Baill. Through the determination of soluble sugar, soluble protein, hormone and peroxidase isoenzyme in the process of formation of rhizome, we try to reveal the relations between the formation mechanism and these factors.The experiment results were as follows:
1. Development of the rhizome was similar to stem, both of them can be divided into promeristem, primary and secondary growth three stages. However, the ultrastructure of primary growth was different, for example, there was a special membrane clinging on the first layer cell of stem tip, and it was much more developed than rhizome tip with completed envelope, more plastids and less vacuoles, while the rhizome tip had more vacuoles and lipid droplets. Furthermore, they had other differences in tip type and secondary structure, for example, the rhizome tip was bendy and the proportion of secondary growth of the rhizome was less than stem.
2. Secondary structure of root was composed of periderm, cambium layer, secondary xylem and phloem, while we didn’t find the parenchyma cells with lignification, developing patterns of primary structure of rhizome is endarch, which is different of developing patterns of primary structure of root.
3. One year axillary bud of cotyledon could develop to rhizome in late August-early September slowly, and the axillary bud of rough leaf to the eighth euphylla would develop into rhizome finally when the condition was appropriate. Rotiform rhizomes were concentrated and whorled in the base of stem. Ultrastructure of axillary bud on the higher node have more differences from rough leaf, it would be decided in early stage that whether axillary bud of cotyledon develop to rhizome or not.
4. Rhizome tip of Schisandra chinensis (Turcz.) Baill had higher content of soluble sugar at initial and dormant period, which reflects the rhizome is different with other plants’rhizome. We found the rhizome not only had the reproductive function, but also had certain of storage feature, so the rhizome competed with the ground on nutrition. There was relationship between the content of soluble protein and the strong level of growth, and in the vigorous growth of rhizome, soluble protein content was consequently higher.
5. There was a great difference of peroxidase isoenzyme map in different developmental period and different type of buds in the process of the formation and development rhizome. The buds which can develop into rhizome have two more POD isoenzyme bands than the buds can’t develop into rhizome. The two POD isoenzyme bands is specific zymogram.
6. Levels of ABA and ZR were higher in formation period of rhizomes of Schisandra chinensis (Turcz.) Baill, which were conducive to induce rhizome. Levels IAA and GA3 were higher in the elongation growth stage, which were essential to rhizome elongation.
引文
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