何首乌和木立芦荟的结构发育与有效成分积累相关性的研究
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摘要
何首乌(Polygonum multiflorum Thunb.)隶属于蓼科(Polygonaceae)蓼属(Polygonum L.),为多年生草质藤本植物。何首乌的主要药用部分是块根,即中药何首乌,以生首乌和制首乌二种形式入药,地上部分的茎叶也可入药,即中药夜交藤。何首乌的主要药效成分是二苯乙烯苷,以及蒽醌类物质。本文运用植物解剖学、组织化学和植物化学等方法,系统研究了何首乌幼苗及成年植株的解剖结构,块根的发生发育规律,蒽醌类物质在何首乌营养器官中的组织化学定位,二苯乙烯苷和蒽醌类物质积累的动态变化规律。
木立芦荟(Aloe arborescens Mill.)隶属于芦荟科(Asphodelaceae)芦荟属(Aloe L.),为多年生肉质草本植物。木立芦荟的主要药用部位是叶,叶内贮藏的芦荟素等蒽醌类物质是其主要药用成分。本文运用石蜡切片、薄切片和超薄切片等植物解剖学方法并结合组织化学、荧光技术和细胞化学等方法,研究木立芦荟叶内蒽醌类物质贮藏部位及贮藏结构的形态本质,并总结出研究蒽醌类物质的细胞化学方法,进而探讨蒽醌类物质的合成场所及运输过程。
通过对何首乌和木立芦荟的结构发育与有效成分积累相关性较为系统的研究,取得了以下主要结果。
何首乌幼苗类型为木兰型。主根四原型,侧根四～六原型,通常无髓部。子叶和初生叶的上表皮无气孔器分布,表皮细胞垂周壁平直;下表皮的表皮细胞垂周壁波浪状,有较多的气孔器,气孔器属无规则型。子叶柄中有3个维管束,初生叶的叶柄中有6个维管束。
何首乌幼苗的过渡区位于下胚轴,根的4个放射维管束,在下胚轴中通过初生韧皮部的拉长、断裂、合拢,形成4个并生维管束,以后相对的2个维管束均一分为二,成为6个维管束。下胚轴在分裂的维管束处裂成两部分,每部分各具3个维管束,与两片子叶的各3个子叶迹相连。根—下胚轴—子叶的维管束成为
Polygonum multiflorum Thunb. (Polygonaceae) is a perennial herbaceous liana. Its root tuber and processed product, namely Chinese traditional medicine He-Shou-Wu, are mainly officinal part. Its stem and leaf, namely Chinese traditional medicine Ye-Jiao-Teng, are also officinal part. The mainly biological activity components of the plant are 2,3,5,4'-tetrahydroxy-stilbene-2-O-β-D- glucoside (for short stilbene glucoside) and anthraquinones. In this study, we systematically investigated the development and structure of the root tuber, the anatomy structure of seedling and adult plant, the histochemistry orientation of anthraquinones in vegetative organ and the accumulation of stilbene glucoside and anthraquinones by using anatomy, histochemistry and phytochemistry methods.Aloe arborescens Mill. (Asphodelaceae) is a perennial succulent herbaceous plant. Anthraquinones are the major medicinal components which are extracted from its leaves. In this study, we investigated the store position of anthraquinones in leaves and the morphological essence of store structure, groped for a cytochemistry method studying anthraquinones, and probed into the synthesize position and transport process. We used the multiple approaches-anatomy, histochemistry, cytochemistry and fluorescence microscopy methods.In this paper, we systematically studied structures and their correlation to the accumulation of effectual component in Polygonum multiflorum and Aloe arborescens, and some important results were listed hereinafter.The seedling type of Polygonum multiflorum is Magnolia Type. Primary xylem in main root is tetrarch, and in lateral root are tetrarch, pentarch and hexarch. The
adaxial epidermis of cotyledon and primary leaf hasn't stomatal apparatus, the anticlinal wall of epidermis cell is straight. The anticlinal wall of epidermis cell is wavy, and many stomatal apparatus which is anomocytic type are distributed in abaxial epidermis of cotyledon and primary leaf. There are three vascular bundles in the petiole of cotyledon, and six vascular bundles in the petiole of primary leaf.The seedling's transition region of Polygonum multiflorum lies in its hypocotyl. Four radiate bundles of the root come into being four collateral bundles via elongating, rupturing and folding of primary phloem in the hypocotyl. Every two opposite vascular bundles among four collateral bundles will divide into two parts, thus there are six vascular bundles in the hypocotyl. The hypocotyl separate into two parts from partite vascular bundles, there are three vascular bundles in every part, and the three vascular bundles join with three traces of each cotyledon. The vascular bundles of root, hypocotyl and cotyledon turn into primary vascular system, whereas the vascular bundle from the stem comes into being alone.The root tip of Polygonum multiflorum is composed of root cap, meristematic zone, elongation zone and maturation zone. The primary structures of the root and the stem consist of epidermis, cortex and vascular cylinder. Some cells in periderm and phloem parenchyma show red brown, and indicate that these cells contain much anthraqumones. There are two vascular bundles in main vein, its xylem is downwards and phloem upwards in upper vascular bundle, but the other way round in below vascular bundle. There are six vascular bundles among which five small vascular bundles arrange in one ring and the other large one lays centrally in the petiole. Outer flower's tepal extends to a calcar. There are four pollen sac in mature anther, and much papillary above filament. The ovary has a locule, and an orthotropous ovule. The achene is trilateral, and has abundant endosperm.The root tuber of Polygonum multiflorum derives accessory cambium from phloem parenchyma after forming ecumenical secondary structure. The accessory cambium forms tertiary xylem inside, and tertiary phloem outside. Abnormal vascular bundles are composed of the tertiary phloem, accessory cambium and the tertiary xylem, and belong to amphicribral bundle. Some phloem parenchyma cells among
abnormal vascular bundles have been formed can resume dividing and form small abnormal vascular bundles. These abnormal cambiums produce a lot of phloem parenchyma cells, which results in continuous augmenting of the root tuber. The root tuber has 4~8 protrudent pointedness, and present cloudy pattern, because of 4~8 abnormal vascular bundles formed in cross section.Histochemistry test in the vegetation organs showed that the content of anthraquinones in the root tuber is higher than that in the stem and leaf. The periderm and phloem of the root tuber present carmine, these two parts are provided with much anthraquinones. But the xylem doesn't, showing no anthraquinones in the xylem. The periderm and phloem of the stem present fleet red, they contain a little anthraquinones. The epidermis, mesophyll and phloem of the leaf present fleet red, but the xylem doesn't. The phloems in the stem and leaf show color but the xylem doesn't, which reveal that anthraquinones transfer through the phloem.The content of stilbene glucoside in the root tuber is the highest in vegetative organs of Polygonum multiflorum, but in the young parts of vegetative organs and generative organ it is low. It is gradually increased in the root tuber and old stem with adding of growth years, its change in the stem and the leaf are not obvious. It increase from June to October, reach the highest value in October, then drop gradually. It is the highest in periderm in parts of the root tuber, lowest in the phloem. Its difference is much great in the root tuber and its processed product from different areas. The content of anthraquinones is the highest in the root tuber, and low in the other parts. It is gradually increased in the root tuber with adding of growth years. It is high in periderm and phloem and low in xylem in the root tuber.The leaf of Aloe arborescens consists of epidermis, assimilating tissue, aqueous tissue and vascular bundles. Vascular bundles, which are composed of one layer of parenchyma cells as a vascular bundle sheath, phloem and xylem tissues, are located between the assimilating tissue and aqueous tissues. There are several large parenchyma cells, i.e. aloin cells, at the phloem pole of each bundle. The aloin cells occupy the most of the area of the cross section of a bundle. The vascular bundle sheath cell stemmed from the ground meristem, its morphological essential is
parenchyma tissue. But these aloin cells come of a procambial cell outside the sieve tube of the protophloem, it is a special phloem parenchyma cell.The aloin cells produce special precipitation when treated with 5% Pb(CH3COOH)2 solution and show red colour reaction when treated with 5% NaOH solution, but the precipitation and the colour reaction do not appeared in other parts. The unique yellow fluorescence of anthraquinones is found in aloin cells under the fluorescence microscope. The histochemical and fluorescent tests indicated that the aloin cells are the store location of anthraquinones.We found an ultracytochemical localization method of aloin, the lead acetate precipitation method, by which the processes of aloin production, transport and storage are observed in transmission electron microscope. Results showed that the thylakoid of plastids come into being aloin precipitation along with vacuolization of assimilating tissue in developing course of Aloe arborescens leave. The amount of aloin precipitation gradually increases with development of the assimilating tissue. There is a lot of dark aloin precipitation in the thylakoid of mature plastids, but other parts of cells are not. So plastid of the assimilating tissue is synthetical location of the aloin. The aloin is transported through the plastid membrane to the surrounding endoplasmic reticulum and enveloped in the vesicles by the endoplasmic reticulum elements, the vesicles approach and later fuse with the plasmalemma. Some vesicles of the plastid membrane directly fuse with the plasmalemma. The vesicles release their contents into the apoplast through exocytosis, and finally reach the vascular bundle sheath by apoplastic translocation. The aloin is transported to the internal tangential wall of vascular bundle sheath cell through endoplasmic reticulum vesicles, and reaches the cytoplasm of the aloin cell by means of plasmodesmata. Finally, the aloin is stored in the vacuoles of the aloin cell.
木立芦荟(Aloe arborescens Mill.)隶属于芦荟科(Asphodelaceae)芦荟属(Aloe L.),为多年生肉质草本植物。木立芦荟的主要药用部位是叶,叶内贮藏的芦荟素等蒽醌类物质是其主要药用成分。本文运用石蜡切片、薄切片和超薄切片等植物解剖学方法并结合组织化学、荧光技术和细胞化学等方法,研究木立芦荟叶内蒽醌类物质贮藏部位及贮藏结构的形态本质,并总结出研究蒽醌类物质的细胞化学方法,进而探讨蒽醌类物质的合成场所及运输过程。
通过对何首乌和木立芦荟的结构发育与有效成分积累相关性较为系统的研究,取得了以下主要结果。
何首乌幼苗类型为木兰型。主根四原型,侧根四～六原型,通常无髓部。子叶和初生叶的上表皮无气孔器分布,表皮细胞垂周壁平直;下表皮的表皮细胞垂周壁波浪状,有较多的气孔器,气孔器属无规则型。子叶柄中有3个维管束,初生叶的叶柄中有6个维管束。
何首乌幼苗的过渡区位于下胚轴,根的4个放射维管束,在下胚轴中通过初生韧皮部的拉长、断裂、合拢,形成4个并生维管束,以后相对的2个维管束均一分为二,成为6个维管束。下胚轴在分裂的维管束处裂成两部分,每部分各具3个维管束,与两片子叶的各3个子叶迹相连。根—下胚轴—子叶的维管束成为
Polygonum multiflorum Thunb. (Polygonaceae) is a perennial herbaceous liana. Its root tuber and processed product, namely Chinese traditional medicine He-Shou-Wu, are mainly officinal part. Its stem and leaf, namely Chinese traditional medicine Ye-Jiao-Teng, are also officinal part. The mainly biological activity components of the plant are 2,3,5,4'-tetrahydroxy-stilbene-2-O-β-D- glucoside (for short stilbene glucoside) and anthraquinones. In this study, we systematically investigated the development and structure of the root tuber, the anatomy structure of seedling and adult plant, the histochemistry orientation of anthraquinones in vegetative organ and the accumulation of stilbene glucoside and anthraquinones by using anatomy, histochemistry and phytochemistry methods.Aloe arborescens Mill. (Asphodelaceae) is a perennial succulent herbaceous plant. Anthraquinones are the major medicinal components which are extracted from its leaves. In this study, we investigated the store position of anthraquinones in leaves and the morphological essence of store structure, groped for a cytochemistry method studying anthraquinones, and probed into the synthesize position and transport process. We used the multiple approaches-anatomy, histochemistry, cytochemistry and fluorescence microscopy methods.In this paper, we systematically studied structures and their correlation to the accumulation of effectual component in Polygonum multiflorum and Aloe arborescens, and some important results were listed hereinafter.The seedling type of Polygonum multiflorum is Magnolia Type. Primary xylem in main root is tetrarch, and in lateral root are tetrarch, pentarch and hexarch. The
adaxial epidermis of cotyledon and primary leaf hasn't stomatal apparatus, the anticlinal wall of epidermis cell is straight. The anticlinal wall of epidermis cell is wavy, and many stomatal apparatus which is anomocytic type are distributed in abaxial epidermis of cotyledon and primary leaf. There are three vascular bundles in the petiole of cotyledon, and six vascular bundles in the petiole of primary leaf.The seedling's transition region of Polygonum multiflorum lies in its hypocotyl. Four radiate bundles of the root come into being four collateral bundles via elongating, rupturing and folding of primary phloem in the hypocotyl. Every two opposite vascular bundles among four collateral bundles will divide into two parts, thus there are six vascular bundles in the hypocotyl. The hypocotyl separate into two parts from partite vascular bundles, there are three vascular bundles in every part, and the three vascular bundles join with three traces of each cotyledon. The vascular bundles of root, hypocotyl and cotyledon turn into primary vascular system, whereas the vascular bundle from the stem comes into being alone.The root tip of Polygonum multiflorum is composed of root cap, meristematic zone, elongation zone and maturation zone. The primary structures of the root and the stem consist of epidermis, cortex and vascular cylinder. Some cells in periderm and phloem parenchyma show red brown, and indicate that these cells contain much anthraqumones. There are two vascular bundles in main vein, its xylem is downwards and phloem upwards in upper vascular bundle, but the other way round in below vascular bundle. There are six vascular bundles among which five small vascular bundles arrange in one ring and the other large one lays centrally in the petiole. Outer flower's tepal extends to a calcar. There are four pollen sac in mature anther, and much papillary above filament. The ovary has a locule, and an orthotropous ovule. The achene is trilateral, and has abundant endosperm.The root tuber of Polygonum multiflorum derives accessory cambium from phloem parenchyma after forming ecumenical secondary structure. The accessory cambium forms tertiary xylem inside, and tertiary phloem outside. Abnormal vascular bundles are composed of the tertiary phloem, accessory cambium and the tertiary xylem, and belong to amphicribral bundle. Some phloem parenchyma cells among
abnormal vascular bundles have been formed can resume dividing and form small abnormal vascular bundles. These abnormal cambiums produce a lot of phloem parenchyma cells, which results in continuous augmenting of the root tuber. The root tuber has 4~8 protrudent pointedness, and present cloudy pattern, because of 4~8 abnormal vascular bundles formed in cross section.Histochemistry test in the vegetation organs showed that the content of anthraquinones in the root tuber is higher than that in the stem and leaf. The periderm and phloem of the root tuber present carmine, these two parts are provided with much anthraquinones. But the xylem doesn't, showing no anthraquinones in the xylem. The periderm and phloem of the stem present fleet red, they contain a little anthraquinones. The epidermis, mesophyll and phloem of the leaf present fleet red, but the xylem doesn't. The phloems in the stem and leaf show color but the xylem doesn't, which reveal that anthraquinones transfer through the phloem.The content of stilbene glucoside in the root tuber is the highest in vegetative organs of Polygonum multiflorum, but in the young parts of vegetative organs and generative organ it is low. It is gradually increased in the root tuber and old stem with adding of growth years, its change in the stem and the leaf are not obvious. It increase from June to October, reach the highest value in October, then drop gradually. It is the highest in periderm in parts of the root tuber, lowest in the phloem. Its difference is much great in the root tuber and its processed product from different areas. The content of anthraquinones is the highest in the root tuber, and low in the other parts. It is gradually increased in the root tuber with adding of growth years. It is high in periderm and phloem and low in xylem in the root tuber.The leaf of Aloe arborescens consists of epidermis, assimilating tissue, aqueous tissue and vascular bundles. Vascular bundles, which are composed of one layer of parenchyma cells as a vascular bundle sheath, phloem and xylem tissues, are located between the assimilating tissue and aqueous tissues. There are several large parenchyma cells, i.e. aloin cells, at the phloem pole of each bundle. The aloin cells occupy the most of the area of the cross section of a bundle. The vascular bundle sheath cell stemmed from the ground meristem, its morphological essential is
parenchyma tissue. But these aloin cells come of a procambial cell outside the sieve tube of the protophloem, it is a special phloem parenchyma cell.The aloin cells produce special precipitation when treated with 5% Pb(CH3COOH)2 solution and show red colour reaction when treated with 5% NaOH solution, but the precipitation and the colour reaction do not appeared in other parts. The unique yellow fluorescence of anthraquinones is found in aloin cells under the fluorescence microscope. The histochemical and fluorescent tests indicated that the aloin cells are the store location of anthraquinones.We found an ultracytochemical localization method of aloin, the lead acetate precipitation method, by which the processes of aloin production, transport and storage are observed in transmission electron microscope. Results showed that the thylakoid of plastids come into being aloin precipitation along with vacuolization of assimilating tissue in developing course of Aloe arborescens leave. The amount of aloin precipitation gradually increases with development of the assimilating tissue. There is a lot of dark aloin precipitation in the thylakoid of mature plastids, but other parts of cells are not. So plastid of the assimilating tissue is synthetical location of the aloin. The aloin is transported through the plastid membrane to the surrounding endoplasmic reticulum and enveloped in the vesicles by the endoplasmic reticulum elements, the vesicles approach and later fuse with the plasmalemma. Some vesicles of the plastid membrane directly fuse with the plasmalemma. The vesicles release their contents into the apoplast through exocytosis, and finally reach the vascular bundle sheath by apoplastic translocation. The aloin is transported to the internal tangential wall of vascular bundle sheath cell through endoplasmic reticulum vesicles, and reaches the cytoplasm of the aloin cell by means of plasmodesmata. Finally, the aloin is stored in the vacuoles of the aloin cell.
引文
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