曼陀罗不同居群形态多样性及生物碱含量比较研究
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摘要
曼陀罗(Datura stramonium L.)又称醉心花、万桃花、狗核桃、洋金花,具有久远的药用历史,还可作观赏植物。国内各省区均有分布,多生于田边、路边及宅旁等地,较少栽培,大多野生。我国地大物博,不同生长地域环境的差异,使曼陀罗具有了丰富的形态多样性。为了更好保护和合理开发利用曼陀罗资源,我们对分布于国内陕西、河北、辽宁、山东等地的曼陀罗10个居群的形态及生物碱的多样性进行了初步比较研究。
以集中种植在杨凌的10个不同居群曼陀罗为研究对象,采用植株形态比较、叶表皮微形态特征分析和种子、茎、叶不同器官生物碱(莨菪碱、阿托品)含量测定,比较了10个居群曼陀罗的形态多样性和生物碱含量差异,得出主要结论如下:
1.种植在同一地的曼陀罗不同居群间的物候期有差异。陕西渭南和陕西商洛的出苗较早,开花较早,衰亡较早,生活期较短。辽宁铁岭的出苗晚,开花早,营养生长期较短,而生殖生长期较长。河北保定的出苗晚,开花也较晚,营养生长期较辽宁铁岭营养生长期长。其他居群的生活期在陕西商洛、陕西渭南和河北保定、辽宁铁岭的居群之间。
2.曼陀罗不同居群抗逆性有差异。河北保定曼陀罗黑斑病和叶片皱缩程度最轻,陕西凤县和辽宁铁岭叶片皱缩最严重;杨凌水运曼陀罗居群黑斑病现象最严重。花色为紫色的曼陀罗居群的抗逆性高于花色为白色曼陀罗居群。
3.不同居群曼陀罗的种子表面纹饰差异主要表现在:(1)种子表面的凹穴排列方式;(2)种脐相对位置;(3)种脐形状等的差异。
4.曼陀罗种内居群的形态变异丰富,变异系数变化范围为4.64%～62.24%。居群间叶、茎、花等形态性状均存在显著差异,其中营养器官的形态较繁殖器官具有更大的遗传变异,部分繁殖器官特性(如鲜果十粒重)在居群间变异也较大。
5.曼陀罗形态性状数据聚类分析可将10个居群分为三类,杨凌水运与辽宁铁岭聚为一类;河北保定、陕西渭南、陕西临潼聚为一类;杨凌大寨乡、陕西商洛、山东莱阳、陕西榆林、陕西凤县聚为一类。
6.曼陀罗叶表皮细胞形状有无规则型和多边形两种,垂周壁式样为波状,但波的深浅不同,分为深波状和浅波状。
7.曼陀罗叶片的上下表皮都存在气孔,气孔属于无规则型。除山东莱阳居群上表皮气孔指数大于下表皮气孔指数外,其它9个居群叶下表皮的气孔指数、气孔密度均大于上表皮的气孔指数、气孔密度。
8.同一曼陀罗居群的不同器官中主要生物碱含量存在差异。表现为种子和茎中阿托品含量都明显高于莨菪碱含量;叶片中,除河北、商洛居群外,其他8个居群的阿托品含量也都高于莨菪碱含量。在同一居群中总生物碱含量均为种子>叶>茎。
9.不同居群曼陀罗的同一器官中莨菪碱和阿托品含量均存在差异。种子中总生物碱含量依次为:陕西渭南>山东莱阳>杨凌大寨乡>辽宁铁岭>陕西凤县>陕西榆林>陕西商洛>杨凌水运>河北保定>陕西临潼;茎中总生物碱依次为:陕西渭南>杨凌大寨乡>陕西凤县>陕西榆林>陕西临潼>山东莱阳>杨凌水运>河北保定>陕西商洛>辽宁铁岭;叶中总生物碱含量依次为:陕西渭南>陕西商洛>陕西临潼>杨凌大寨乡>陕西榆林>辽宁铁岭>山东莱阳>陕西凤县>河北保定>杨凌水运。
Datura stramonium L. (Jimson weed, thorn apple), which could be also regarded as ornamental plant, had a long story as medicinal plant. D. stramonium L. (mostly wild) distributed in most provinces in China, and was always found alongside fields, roadside, waste land, etc. Geographical environments and climates were quite different around China, so D. stramonium had diversity in morphology. For the purpose of preserve and make use of D. stramonium L. resources, our studies were on the morphology and alkaloid (hyoscyamine, atropine) comparison between different populations from Shaanxi, Hebei, Liaoning, Shandong, etc. 10 different populations were grown in Yangling as the test materials, whose plant morphology indexes, micromorphology of leaf epidermis, alkaloid contents of seeds, leaves, stems were determined. The test results were as follows:
1. Although planted in the same fields, The ghenoghases of D. stramonium L. differed in different populations. Seedling emergences and initial bloom of populations from Weinan, Shaanxi and Shangluo, Shaanxi were early, so they were decline phase, thus their ghenoghases were short; Populations from Tieling, Liaoning had a late seedling emergence and a early blossom, that made it a short vegetative growth and a long reproduction growth; Baoding, Hebei population’s emergence and initial bloom was late, the vegetative growth was longer than that of Tieling, Liaoning population. There were always flowers blooming in D. stramonium L. plant during the reproduction growth. The life phases of the other populations were among populations above.
2. Stress resistances of different populations were different. Population from Baoding, Hebei had the slightest black spot disease and leaf shrinkage. The leaf shrinkage of Fengxian, Shaanxi and Tieling, Liaoning population were the most severe, while Shuiyun, Yangling had a most serious black spot disease. Stress resistances of D. stramonium L. with purple flowers was higher than that with white flowers.
3. The seeds surface ornamentations differences of different D. stramonium L. populations mainly were size and arrangement pattern of seeds surface concave points, relative position of hilums, shape of hilums, etc.
4. there were always morphology variation in different populations of D. stramonium L. species. Variable range of variation coefficients was 4.64%～62.24%. The leaves, stems and flowers’morphology indexes were significantly different within populations. Besides, vegetative organs had more genetic variations than reproductive organs. Nevertheless, some of reproductive organs also had significant variation.
5. The cluster analysis of morphology indexes showed that 10 D. stramonium L. populations could be divided into 3 clusters: Shuiyun, Yangling and Tieling, Liaoning; Baoding, Hebei, Weinan, Shaanxi, and Lintong, Shaanxi; Dazhaixiang, Yangling, Shangluo, Shaanxi, Laiyang, Shandong, Yulin, Shaanxi and Fengxian, Shaanxi.
6. Shape of D. stramonium L.leaf epidermis cells had two different forms: irregular and polygonal. The form of anticlinal walls was wavy, which had various extent: deeply wavy and slightly wavy.
7. There were stomata, whose shape were irregular, on both sides of D. stramonium L. leaf epidermis. Stomatal indexes of lower epidermis of all 10 populations were higher than upper epidermis, except for population from Laiyang, Shandong.
8. In the same D. stramonium L. population, Main alkaloid contents in different organs are varied. In stems and seeds, atropine contents were higher than hyoscyamine contents. As to leaves, except for populations from Hebei and Shangluo, atropine contents of all the populations were higher than hyoscyamine contents. In every single population, the order of alkaloid contents was seeds > leaves > stems.
9.Content difference of hyoscyamine and atropine in the same organic of different populations. Alkaloid Content of seed in order:SW>LL>YL1>LT>SF>SY>SS>YL2>HB>SL; Alkaloid Content of stem in order:SW>YL2>SF>SY>SL>LL>YL2>HB>SS>LT; Alkaloid Content of leaf in order:SW>SS>SL>YL1>SY>LT>LL>SF>HB>YL2.
以集中种植在杨凌的10个不同居群曼陀罗为研究对象,采用植株形态比较、叶表皮微形态特征分析和种子、茎、叶不同器官生物碱(莨菪碱、阿托品)含量测定,比较了10个居群曼陀罗的形态多样性和生物碱含量差异,得出主要结论如下:
1.种植在同一地的曼陀罗不同居群间的物候期有差异。陕西渭南和陕西商洛的出苗较早,开花较早,衰亡较早,生活期较短。辽宁铁岭的出苗晚,开花早,营养生长期较短,而生殖生长期较长。河北保定的出苗晚,开花也较晚,营养生长期较辽宁铁岭营养生长期长。其他居群的生活期在陕西商洛、陕西渭南和河北保定、辽宁铁岭的居群之间。
2.曼陀罗不同居群抗逆性有差异。河北保定曼陀罗黑斑病和叶片皱缩程度最轻,陕西凤县和辽宁铁岭叶片皱缩最严重;杨凌水运曼陀罗居群黑斑病现象最严重。花色为紫色的曼陀罗居群的抗逆性高于花色为白色曼陀罗居群。
3.不同居群曼陀罗的种子表面纹饰差异主要表现在:(1)种子表面的凹穴排列方式;(2)种脐相对位置;(3)种脐形状等的差异。
4.曼陀罗种内居群的形态变异丰富,变异系数变化范围为4.64%～62.24%。居群间叶、茎、花等形态性状均存在显著差异,其中营养器官的形态较繁殖器官具有更大的遗传变异,部分繁殖器官特性(如鲜果十粒重)在居群间变异也较大。
5.曼陀罗形态性状数据聚类分析可将10个居群分为三类,杨凌水运与辽宁铁岭聚为一类;河北保定、陕西渭南、陕西临潼聚为一类;杨凌大寨乡、陕西商洛、山东莱阳、陕西榆林、陕西凤县聚为一类。
6.曼陀罗叶表皮细胞形状有无规则型和多边形两种,垂周壁式样为波状,但波的深浅不同,分为深波状和浅波状。
7.曼陀罗叶片的上下表皮都存在气孔,气孔属于无规则型。除山东莱阳居群上表皮气孔指数大于下表皮气孔指数外,其它9个居群叶下表皮的气孔指数、气孔密度均大于上表皮的气孔指数、气孔密度。
8.同一曼陀罗居群的不同器官中主要生物碱含量存在差异。表现为种子和茎中阿托品含量都明显高于莨菪碱含量;叶片中,除河北、商洛居群外,其他8个居群的阿托品含量也都高于莨菪碱含量。在同一居群中总生物碱含量均为种子>叶>茎。
9.不同居群曼陀罗的同一器官中莨菪碱和阿托品含量均存在差异。种子中总生物碱含量依次为:陕西渭南>山东莱阳>杨凌大寨乡>辽宁铁岭>陕西凤县>陕西榆林>陕西商洛>杨凌水运>河北保定>陕西临潼;茎中总生物碱依次为:陕西渭南>杨凌大寨乡>陕西凤县>陕西榆林>陕西临潼>山东莱阳>杨凌水运>河北保定>陕西商洛>辽宁铁岭;叶中总生物碱含量依次为:陕西渭南>陕西商洛>陕西临潼>杨凌大寨乡>陕西榆林>辽宁铁岭>山东莱阳>陕西凤县>河北保定>杨凌水运。
Datura stramonium L. (Jimson weed, thorn apple), which could be also regarded as ornamental plant, had a long story as medicinal plant. D. stramonium L. (mostly wild) distributed in most provinces in China, and was always found alongside fields, roadside, waste land, etc. Geographical environments and climates were quite different around China, so D. stramonium had diversity in morphology. For the purpose of preserve and make use of D. stramonium L. resources, our studies were on the morphology and alkaloid (hyoscyamine, atropine) comparison between different populations from Shaanxi, Hebei, Liaoning, Shandong, etc. 10 different populations were grown in Yangling as the test materials, whose plant morphology indexes, micromorphology of leaf epidermis, alkaloid contents of seeds, leaves, stems were determined. The test results were as follows:
1. Although planted in the same fields, The ghenoghases of D. stramonium L. differed in different populations. Seedling emergences and initial bloom of populations from Weinan, Shaanxi and Shangluo, Shaanxi were early, so they were decline phase, thus their ghenoghases were short; Populations from Tieling, Liaoning had a late seedling emergence and a early blossom, that made it a short vegetative growth and a long reproduction growth; Baoding, Hebei population’s emergence and initial bloom was late, the vegetative growth was longer than that of Tieling, Liaoning population. There were always flowers blooming in D. stramonium L. plant during the reproduction growth. The life phases of the other populations were among populations above.
2. Stress resistances of different populations were different. Population from Baoding, Hebei had the slightest black spot disease and leaf shrinkage. The leaf shrinkage of Fengxian, Shaanxi and Tieling, Liaoning population were the most severe, while Shuiyun, Yangling had a most serious black spot disease. Stress resistances of D. stramonium L. with purple flowers was higher than that with white flowers.
3. The seeds surface ornamentations differences of different D. stramonium L. populations mainly were size and arrangement pattern of seeds surface concave points, relative position of hilums, shape of hilums, etc.
4. there were always morphology variation in different populations of D. stramonium L. species. Variable range of variation coefficients was 4.64%～62.24%. The leaves, stems and flowers’morphology indexes were significantly different within populations. Besides, vegetative organs had more genetic variations than reproductive organs. Nevertheless, some of reproductive organs also had significant variation.
5. The cluster analysis of morphology indexes showed that 10 D. stramonium L. populations could be divided into 3 clusters: Shuiyun, Yangling and Tieling, Liaoning; Baoding, Hebei, Weinan, Shaanxi, and Lintong, Shaanxi; Dazhaixiang, Yangling, Shangluo, Shaanxi, Laiyang, Shandong, Yulin, Shaanxi and Fengxian, Shaanxi.
6. Shape of D. stramonium L.leaf epidermis cells had two different forms: irregular and polygonal. The form of anticlinal walls was wavy, which had various extent: deeply wavy and slightly wavy.
7. There were stomata, whose shape were irregular, on both sides of D. stramonium L. leaf epidermis. Stomatal indexes of lower epidermis of all 10 populations were higher than upper epidermis, except for population from Laiyang, Shandong.
8. In the same D. stramonium L. population, Main alkaloid contents in different organs are varied. In stems and seeds, atropine contents were higher than hyoscyamine contents. As to leaves, except for populations from Hebei and Shangluo, atropine contents of all the populations were higher than hyoscyamine contents. In every single population, the order of alkaloid contents was seeds > leaves > stems.
9.Content difference of hyoscyamine and atropine in the same organic of different populations. Alkaloid Content of seed in order:SW>LL>YL1>LT>SF>SY>SS>YL2>HB>SL; Alkaloid Content of stem in order:SW>YL2>SF>SY>SL>LL>YL2>HB>SS>LT; Alkaloid Content of leaf in order:SW>SS>SL>YL1>SY>LT>LL>SF>HB>YL2.
引文
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