甘遂乳汁管的分布、发育与超微结构以及6种大戟属植物的比较研究
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摘要
大戟属(Euphorbia L.)植物隶属大戟科(Euphorbiaceae),全世界约有2000种,本属植物中所含化学成分具有明显的生理活性,已有一些种类应用于防病治病。民间用来治疗通便、利尿、治疗水肿、结核、牛皮癣、疥疮和无名肿毒,尤其是除疣、抗肿瘤等症,具有非常重要的药用价值。因此,大戟属植物一直就引起人们的广泛关注,对它的研究也深入到各个领域。甘遂(Euphorbia kansui Liou)为大戟属多年生草本植物,主产于陕西、山西、河南和甘肃等地,为《中华人民共和国药典》(2005版)收录的药用植物。因此本论文首先利用石蜡切片、半薄切片和组织化学方法对甘遂各器官中乳汁管的类型、分布和大小进行了研究,并进一步利用透射电子显微镜研究了甘遂发育中及成熟的无节乳汁管的超微结构;其次,以甘遂、泽漆(E.helioscopia Linn.)、地锦草(E.humifusaWilld.)、大戟(E.pekinensis Rupr.)、湖北大戟(E.hylonoma Hand.-Mazz)及华北大戟(E.lunulata Bunge,Enum.)6种大戟属植物为研究对象,利用石蜡切片、叶片离析法、扫描电镜及傅立叶变换红外光谱扫描法系统比较其各器官的结构、叶表皮形态、果实形状及其表皮的微形态以及傅里叶变换红外光谱,并利用SPSS13.0软件进行聚类分析,探讨6种植物间的亲缘关系,为大戟属植物的分类及生药鉴定等提供可靠依据。
1.甘遂根茎叶的发生过程,与多年生双子叶草本植物的一般发育规律相类似。根初生结构分化后形成的初生木质部脊数有二原型、三原型或四原型;甘遂茎的个体发育包括四个阶段,分别是原分生组织、初生分生组织、初生结构与次生生长阶段:叶片由叶原基经过初生生长形成其成熟结构。
2.甘遂组织化学实验结果显示二萜类化合物在甘遂的营养器官中分布广泛。根中柱鞘和韧皮部的薄壁组织细胞,茎皮层、维管形成层和韧皮部的薄壁组织细胞,叶的叶肉细胞以及叶脉中的厚角组织和薄壁组织细胞的腔中都显示不同程度的红色,而乳汁管中乳汁的显色较深。
3.甘遂乳汁管为无节分枝型,在各器官中主要分布在维管束韧皮部的外侧或周围,此外,在根的中柱鞘薄壁细胞、叶片的叶肉组织、果实的中果皮及胚乳细胞内也有少量乳汁管的分布。甘遂乳汁管的大小在各器官中略有不同,其中茎中直径最大为39.79μm,块根中直径居中为36.90μm,叶肉组织中直径最小,仅有8.94μm;乳汁管的密度在其营养器官中差异较大,依次为叶＞茎＞根＞块根。
4.甘遂乳汁管发育的两个主要特点为:(1)形成大量小液泡,其起源主要为内质网;(2)高尔基体释放大量的高尔基体小泡。成熟乳汁管的原生质体为一电子致密薄层,中央为大液泡,其内充满了乳汁颗粒。
5.比较研究结果表明6种大戟属植物的各器官结构、叶表皮特征、叶片及果实表皮微形态和傅里叶变换红外光谱均存在不同程度的差异。应用SPSS13.0对形态结构性状和红外光谱数据及单独的形态结构性状分别进行了聚类分析,结果将6种大戟属植物聚为2类,地锦草、华北大戟、大戟、湖北大戟与泽漆属于一大类,而甘遂属另一类。其中,地锦草与华北大戟、大戟与湖北大戟表现出较强的相似性,两种植物间具有较近的亲缘关系。
Euphorbia L. belonging to Euphorbiaceae has about 2,000 species all over the world. Some ingredients of the plants have obvious physiological activity, as a result, some species have been used for the prevention and treatment of disease. They are primarily used as traditional Chinese medicines to defecate, increase the discharge of urine and treat oedema, tuberculosis, psoriasis, scabies and unknown tumors. Therefore, Euphorbia L. has been paid more attention and concerning studies have also penetrated into all fields. Euphorbia kansui Liou is a perennial herbaceous species in Euphorbia L.. It is mainly distributed in Shannxi, Shanxi, Henan, Gansu and other provinces in China. It is also a relatively important herb in Euphorbia L.. Therefore firstly, the type, distribution and size of laticifers in all organs of E. kansui were observed under light-microscope by the means of paraffin sectioning, semi-thin sectioning and histochemistry, furthermore the structure of developing and mature nonarticulated laticifers in E. kansui were studied at ultrastructural level with transmission electron microscopy; secondly, comparative studies of E. kansui, E. helioscopia, E. humifusa, E. pekinensis, E. hylonoma and E. lunulata were investigated by the methods of paraffin sectioning, leaf epidermis isolating, scanning electron microscopy (SEM) and Fourier Transform Infrared (FTIR) spectroscopy technique. Furthermore, SPSS13.0 software was used for cluster analysis to discuss the relationship among the six species so as to provide reliable evidence for the classification and identification of crude medicine of Euphorbia L..
1. The developing process of root, stem and leaf of E. kansui were similar to the normal developing principles of other dicotyledons. The primary xylem was diarch, triarch or tetrarch. The development of stem could be divided into four stages, which were promeristem, primary meristem, primary structure and secondary growth stages. The mature structure of leaf was formed after the primary growth of phyllopodium.
2. The histochemical results showed that the diterpene esters were distributed widely in the vegetative organs of E. kansui. The color showed in the cell cavity of laticifers was the darkest.
3. The laticifers in E. kansui is non-articulated branched type. The laticifers were mainly distributed around the phloem of the vascular bundle, they were also distributed in the pericycle of root tuber, the mesophyll tissues of leaf, the mesocarp and endosperm in fruit. The size of laticifers in E. kansui varied slightly in different organs. The laticifers in stem were largest (39.79μm), the laticifers in root tuber were the medium (36.90μm) and the laticifers in mesophyll tissues were the smallest (8.94μm) . While, the density of laticifers varied greatly in different vegetative organs and it is leaf, stem, root and root tuber from high to low.
4. The most important two features in the differentiation of the laticifers in E. kansui were the development of small vacuoles arising from ER and releasing lots of Golgi vesicles by Golgi bodies. The mature laticifers had a thin layer of electron-dense peripheral cytoplasm in which the organelle could not be distinguished and a large central vacuole filled with latex particles.
5. The comparative anatomy results revealed that various degrees of differences existed in the structure of organs, the micromorphology of leaves and fruits, and the FTIR spectra. Cluster analyses on the structural characteristics and FTIR data together and the structural characteristics alone were carried out by using SPSS13.0 software respectively. The results showed that six species of Euphorbia can be divided into two clusters. E. humifusa, E. lunulata, E. pekinensis, E. hylonama and E. helioscopia fell into one cluster, while E. kansui fell into the other one. Stronger similarities and relatively close relationships were shown between E. humifusa and E. lunulata as well as between E. pekinensis and E. hylonama.
1.甘遂根茎叶的发生过程,与多年生双子叶草本植物的一般发育规律相类似。根初生结构分化后形成的初生木质部脊数有二原型、三原型或四原型;甘遂茎的个体发育包括四个阶段,分别是原分生组织、初生分生组织、初生结构与次生生长阶段:叶片由叶原基经过初生生长形成其成熟结构。
2.甘遂组织化学实验结果显示二萜类化合物在甘遂的营养器官中分布广泛。根中柱鞘和韧皮部的薄壁组织细胞,茎皮层、维管形成层和韧皮部的薄壁组织细胞,叶的叶肉细胞以及叶脉中的厚角组织和薄壁组织细胞的腔中都显示不同程度的红色,而乳汁管中乳汁的显色较深。
3.甘遂乳汁管为无节分枝型,在各器官中主要分布在维管束韧皮部的外侧或周围,此外,在根的中柱鞘薄壁细胞、叶片的叶肉组织、果实的中果皮及胚乳细胞内也有少量乳汁管的分布。甘遂乳汁管的大小在各器官中略有不同,其中茎中直径最大为39.79μm,块根中直径居中为36.90μm,叶肉组织中直径最小,仅有8.94μm;乳汁管的密度在其营养器官中差异较大,依次为叶＞茎＞根＞块根。
4.甘遂乳汁管发育的两个主要特点为:(1)形成大量小液泡,其起源主要为内质网;(2)高尔基体释放大量的高尔基体小泡。成熟乳汁管的原生质体为一电子致密薄层,中央为大液泡,其内充满了乳汁颗粒。
5.比较研究结果表明6种大戟属植物的各器官结构、叶表皮特征、叶片及果实表皮微形态和傅里叶变换红外光谱均存在不同程度的差异。应用SPSS13.0对形态结构性状和红外光谱数据及单独的形态结构性状分别进行了聚类分析,结果将6种大戟属植物聚为2类,地锦草、华北大戟、大戟、湖北大戟与泽漆属于一大类,而甘遂属另一类。其中,地锦草与华北大戟、大戟与湖北大戟表现出较强的相似性,两种植物间具有较近的亲缘关系。
Euphorbia L. belonging to Euphorbiaceae has about 2,000 species all over the world. Some ingredients of the plants have obvious physiological activity, as a result, some species have been used for the prevention and treatment of disease. They are primarily used as traditional Chinese medicines to defecate, increase the discharge of urine and treat oedema, tuberculosis, psoriasis, scabies and unknown tumors. Therefore, Euphorbia L. has been paid more attention and concerning studies have also penetrated into all fields. Euphorbia kansui Liou is a perennial herbaceous species in Euphorbia L.. It is mainly distributed in Shannxi, Shanxi, Henan, Gansu and other provinces in China. It is also a relatively important herb in Euphorbia L.. Therefore firstly, the type, distribution and size of laticifers in all organs of E. kansui were observed under light-microscope by the means of paraffin sectioning, semi-thin sectioning and histochemistry, furthermore the structure of developing and mature nonarticulated laticifers in E. kansui were studied at ultrastructural level with transmission electron microscopy; secondly, comparative studies of E. kansui, E. helioscopia, E. humifusa, E. pekinensis, E. hylonoma and E. lunulata were investigated by the methods of paraffin sectioning, leaf epidermis isolating, scanning electron microscopy (SEM) and Fourier Transform Infrared (FTIR) spectroscopy technique. Furthermore, SPSS13.0 software was used for cluster analysis to discuss the relationship among the six species so as to provide reliable evidence for the classification and identification of crude medicine of Euphorbia L..
1. The developing process of root, stem and leaf of E. kansui were similar to the normal developing principles of other dicotyledons. The primary xylem was diarch, triarch or tetrarch. The development of stem could be divided into four stages, which were promeristem, primary meristem, primary structure and secondary growth stages. The mature structure of leaf was formed after the primary growth of phyllopodium.
2. The histochemical results showed that the diterpene esters were distributed widely in the vegetative organs of E. kansui. The color showed in the cell cavity of laticifers was the darkest.
3. The laticifers in E. kansui is non-articulated branched type. The laticifers were mainly distributed around the phloem of the vascular bundle, they were also distributed in the pericycle of root tuber, the mesophyll tissues of leaf, the mesocarp and endosperm in fruit. The size of laticifers in E. kansui varied slightly in different organs. The laticifers in stem were largest (39.79μm), the laticifers in root tuber were the medium (36.90μm) and the laticifers in mesophyll tissues were the smallest (8.94μm) . While, the density of laticifers varied greatly in different vegetative organs and it is leaf, stem, root and root tuber from high to low.
4. The most important two features in the differentiation of the laticifers in E. kansui were the development of small vacuoles arising from ER and releasing lots of Golgi vesicles by Golgi bodies. The mature laticifers had a thin layer of electron-dense peripheral cytoplasm in which the organelle could not be distinguished and a large central vacuole filled with latex particles.
5. The comparative anatomy results revealed that various degrees of differences existed in the structure of organs, the micromorphology of leaves and fruits, and the FTIR spectra. Cluster analyses on the structural characteristics and FTIR data together and the structural characteristics alone were carried out by using SPSS13.0 software respectively. The results showed that six species of Euphorbia can be divided into two clusters. E. humifusa, E. lunulata, E. pekinensis, E. hylonama and E. helioscopia fell into one cluster, while E. kansui fell into the other one. Stronger similarities and relatively close relationships were shown between E. humifusa and E. lunulata as well as between E. pekinensis and E. hylonama.
引文
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