香薷叶表皮腺毛及其分泌黄酮类物质的组织化学研究

利用光学显微技术和扫描电镜技术,研究了香薷营养生长期和生殖生长期的枝条上部、中部、下部叶片的腺毛种类、分布及数量;采用组织化学染色和荧光显微技术,观察香薷腺毛分泌黄酮类物质的过程,探讨香薷叶表皮腺毛的泌香机理,为生产中确定提取香薷黄酮类物质的最佳取材时期提供依据。结果表明:(1)香薷叶表皮有头状腺毛和盾状腺毛,成熟的头状腺毛包括基细胞、柄细胞和头部,头部较小,呈半圆球型,直径为(20±2)μm;发育近成熟的盾状腺毛头部较大,呈盾状,直径为(60±5)μm。(2)香薷叶片远轴面和近轴面均有腺毛的分布,远轴面分布较多,近轴面分布较少;盾状腺毛主要分布在叶的远轴面,头状腺毛在叶的两面均有分布;腺毛密度随节位的降低而减少,节位相同时,营养生长期叶片上的腺毛密度(283.9个/mm2)高于生殖生长期(194.4个/mm2)。(3)香薷头状腺毛和盾状腺毛均能分泌黄酮类物质,且随着腺毛的发育成熟,黄酮类物质逐渐积累于腺毛的头部。     

龙葵腺毛发育的细胞学研究

利用光学显微镜、扫描电镜和透射电镜技术,观察了龙葵“四叶一心”期时叶片及茎表皮的腺毛的种类、分布,探究了不同类型腺毛的起源、生长、成熟、分泌、衰老等发育过程的细胞学特征;通过组织化学染色和荧光显微技术,观察了龙葵腺毛成分、分布,为龙葵的进一步开发利用提供参考。结果表明：（1）龙葵腺毛分为单细胞头腺毛和多细胞头腺毛两类,前者主要分布于茎表面和叶上下表皮,后者主要分布于茎表面的单细胞头腺毛之间、叶脉及叶边缘;（2）龙葵腺毛发育起始于表皮细胞突起,单细胞头腺毛行顶端生长,具1-4个柄细胞,四种类型;多细胞头腺毛可再分为一层、两层与三层多细胞头腺毛,另具三种特殊类型;（3）龙葵成熟腺毛具分泌能力,通过皮下空间的物质积累导致腺毛头细胞表面形成突起、包块、破口,最终释放分泌物;而头细胞与柄细胞随即皱缩、衰老。（4）超微结构显示,腺毛头细胞中内质网与高尔基体极为丰富,合成代谢及分泌活动活跃,产生大量包裹嗜锇物质的囊泡,囊泡与细胞壁融合,进而将嗜锇物质转移至细胞壁并积累,随后储存在角质层下的皮下空间直至分泌释放;（5）组织化学染色结果表明,腺毛含有萜类、生物碱、脂类、蛋白质、酚类和多糖。头细胞中主要含有萜类、生物碱、脂类、蛋白质、酚类和中性多糖;柄细胞中主要含有萜类、生物碱、脂类。     

狭叶薰衣草腺毛发育过程中的钙分布研究

采用焦锑酸钾沉淀法测定狭叶薰衣草腺毛发育过程中的钙分布动态,结果表明,腺毛钙的分布表现出了一种特殊的时、空变化过程。腺毛发育早期,腺毛原始细胞中有大量的钙颗粒分布,尤其是在细胞壁、质膜和液泡膜上;到2-细胞与3-细胞时期,钙颗粒的数量明显减少,但颗粒较大,且主要集中在细胞壁、液泡、线粒体上;当腺毛成熟并开始分泌时,角质层下间隙中靠细胞壁上的钙颗粒非常丰富,但基细胞和柄细胞的钙颗粒明显下降,特别是头部细胞的细胞质和液泡中;最后,到分泌后期,角质层间隙中的钙颗粒减少,且不规则地分布在细胞质中。推测在腺毛分泌时期,钙呈现出顶部集中分布的特征,这很可能是分泌的一种信号表现。     

光果甘草营养器官结构及其总黄酮的组织化学定位和含量研究

应用植物解剖学、组织化学和植物化学方法,对光果甘草各营养器官的结构、总黄酮的组织化学定位和含量差异进行了研究.结果显示:(1)光果甘草叶为异面叶,由表皮、叶肉和叶脉组成.叶表皮具腺毛,叶肉中具胶囊细胞,主脉发达;茎由表皮(周皮)、皮层、维管柱组成,其髓中具有粘液细胞;根由周皮、次生维管组织组成,周皮具厚木栓层,次生维管组织中次生木质部和纤维发达.(2)黄酮类物质在叶中分布在表皮、腺毛、胶囊细胞、厚角组织和韧皮部和木质部中的薄壁细胞中;茎中分布在周皮、韧皮部和粘液细胞中;在根中则分布在周皮中.(3)不同营养器官中黄酮类物质含量存在差异:叶＞根茎＞主根＞茎.(4)温度的下降促使黄酮类物质从地上合成器官向地下储藏器官的转运.建议每年可在果熟期和枯萎期之间采挖药材,地上部分收割也作药用,综合利用光果甘草资源.     

细叶益母草叶表面腺毛多样性及发育形态学研究

本文研究了分布在细叶益母草(Leonurus sibiricus)叶表面三种腺毛的发育过程,在此基础上,对2-细胞头状腺毛、4-细胞头状腺毛和8-细胞盾状腺毛的多样性特征进行了讨论。     

冬凌草腺毛的形态学及组织化学研究

利用光学显微镜对药用植物冬凌草地上部分腺毛的形态、分布和组织化学进行了研究。结果表明:(1)冬凌草的叶表皮有3种形态显著不同的毛,即非腺毛、盾状腺毛和头状腺毛;盾状腺毛和头状腺毛均具1个基细胞、1个柄细胞和头部;成熟的盾状腺毛的头部一般由4个分泌细胞组成,而头状腺毛头部由2个分泌细胞组成。(2)组织化学鉴定结果显示:2种腺毛中均含有黄酮类成分,盾状腺毛中还含有单萜、倍半萜等萜类成分;冬凌草甲素可能只存在于盾状腺毛中,但需要更直接的证据证明。研究认为,高密度的盾状腺毛可以作为筛选冬凌草高甲素含量品种的一项重要依据。     

细叶益母草(Leonurus sibiricus)叶表面腺毛多样性及发育形态学研究

本文研究了分布在细叶益母草（Ｌｅｏｎｕｒｕｓｓｉｂｉｒｉｃｕｓ） 叶表面三种腺毛的发育过程,在此基础上,对２细胞头状腺毛、４细胞头状腺毛和８细胞盾状腺毛的多样性特征进行了讨论     

细叶益母草（Leonurus sibiricus）叶表面腺毛多样性及发育形态学 …

本文研究了分布在细叶益母草（Ｌｅｏｎｕｒｕｓ ｓｉｂｉｒｉｃｕｓ）叶表面三种腺毛的发育过程,在此基础上,对２－细胞头状腺毛、４－细胞头状腺毛和８－细胞盾状腺毛的多样性特征进行了讨论。     

木香薷腺毛形态结构发生发育规律的研究

采用常规石蜡切片法及扫描电镜技术对木香薷(Elsholtzia stauntoni Benth)腺毛发生发育及其规律进行了研究。结果表明：木香薷表皮上主要有两种表皮毛：无分泌细胞的表皮毛与有分泌细胞的腺毛。前者包括单细胞乳头状毛、2~3细胞管状毛、分枝状毛及多细胞管状毛;后者包括头状腺毛与盾状腺毛。成熟头状腺毛头部由1、2或4个分泌细胞构成,头部呈圆球形或半圆球形;成熟盾状腺毛头部由8~12个分泌细胞构成,分泌细胞横向扩展形成盾状头部。木香薷腺毛主要在茎端幼叶处大量发生,从茎端第一对幼叶处开始产生;从幼叶期到成熟期均有腺毛发生,大部分腺毛在幼叶期发生发育,只有极少部分在叶的成熟期进行发生发育。     

野薄荷叶腺毛的发育形态学研究

研究了分布于我国东北部的野薄荷叶表面腺毛的结构和发育。主要腺毛有两种：多细胞盾状腺毛和弯曲单细胞头状腺毛。多细胞盾状腺毛由１个基细胞,１个柄细胞和一个由８－１２个分泌细胞构成的头部组成。单细胞头状腺毛由１个基细胞,１个柄细胞和一个分泌细胞头部构成。     

Light, conventional and environmental scanning electron microscopy of the trichomes of Cucurbita pepo subsp. pepo var. styriaca and histochemistry of glandular secretory products

BACKGROUND AND AIMS: In the present study, the differences between glandular and non-glandular trichomes, the secretory process and the method of secretion were studied. Previous studies on leaves of Styrian oil pumpkin (Cucurbia pepo var. styriaca) plants have shown that four morphologically and ontogenetically independent glandular and non-glandular trichome types and one bristle hair type can be distinguished. The four types of trichomes can be categorized into three glandular trichome types: type I, a short-stalked trichome with four head cells including a 'middle-cell', two stalk cells and one basal cell; type II, a long-stalked trichome with two head cells, a 'neck-cell' region and a long stalk area; type IV, a 'stipitate-capitate' trichome with a mesophyll cell basement, a short stalk and a multicellular head; type III, a non-glandular 'columnar-digit' trichome, which consists of two head cells continuous with three-celled stalk, and the basal cell. METHODS: The histochemical studies (the main classes of metabolite in secreted material of glandular trichomes) were conducted in fresh and fixed hand sections, using the following tests: Sudan black B, Nile blue A, osmium tetroxide, neutral red, Naturstoffreagent A, FSA (fuchsin-safranin-astra blue), NADI (naphthol + dimethylparaphenylenediamine) and ruthenium red. Each suggested differences in the intercalations during the ontogenetical development of each trichome during the development stage. KEY RESULTS: The histochemical reactions revealed the main components of the materials secreted by all types of trichomes, which include lipids, flavones and terpenes and the different cell wall compositions. Glandular secretions were observed during environmental scanning electron microscopy (ESEM) and the trichomes compared with those seen by conventional scanning electron microscopy (CSEM). CONCLUSIONS: Scanning electron microscopy and histochemical analysis demonstrated that each of the trichomes studied produced and released secretory products in a characteristic way.     

Ficus carica L. leaf anatomy: Trichomes and solid inclusions

Ficus carica L., a typical plant of the Mediterranean environment, presents leaves covered by an extensive indumentum, and a mesophyll full of solid inclusions. The morphology and ultrastructure of the trichomes, calcium carbonate cystoliths and silicified structures of leaves of F. carica cv Dottato were investigated with light, confocal, scanning electron microscopy, and energy-dispersive X-ray spectroscopy. At the same time, histochemical reactions were also employed to analyse the indumentum composed by glandular and non-glandular trichomes by applying chemical reagents and fluorescence microscopy. Non-glandular and glandular trichomes, capitate, are described. Non-glandular trichomes are unicellular simple, spine-like and present different morphology and sizes. The capitate glandular trichomes are present on leaf adaxial and abaxial surface and consist of one-celled stalk and 3/4 cells spherical head. Histochemical characterisation of leaf hairs revealed the presence of flavonoids, while glandular trichome head cells showed a complex mixture of alkaloids, essential oil and flavonoids. Cu and Al were found in the constitutive structures, spike and dome, of the cystoliths. Several epidermal cells and non-glandular trichomes were silicified. Leaf hairs, trichomes secretions, solid inclusions and silicification of F. carica leaf have significant roles to play in relation to leaf protection from external factors, including high-intensity radiation, herbivores or pathogens.     

线纹香茶菜叶上腺毛发育的细胞学研究

为进行中药溪黄草基原植物的品种鉴定,采用光镜和电镜对线纹香茶菜(原变种)[Isodon lophanthoides var.lophanthoides]叶上腺毛的发育进行细胞学研究。结果表明,线纹香茶菜具有头状腺毛和盾状腺毛2种类型。头状腺毛无色透明,由1个基细胞、1个柄细胞和1或2个头部分泌细胞构成;盾状腺毛为红色,由1或2个基细胞、1个柄细胞和4~8个分泌细胞构成头部。2种腺毛均由原表皮细胞经两次平周分裂形成,后因柄细胞和头部细胞所处的分化状态不同而形成两类腺毛。2种腺毛超微结构表明,质体、高尔基体和粗面内质网为主要分泌物产生和运输的细胞器。当盾状腺毛成熟时,角质层下间隙充满了分泌物,其分泌物的性质很可能决定了线纹香茶菜腺毛的颜色。     

Development and ultrastructure of glandular trichomes in<Emphasis Type="Italic">pelargonium x fragrans</Emphasis> ‘mabel grey’ (geraniaceae)

The glandular trichomes of leaves fromPelargonium xfragrans ‘Mabel Grey’ (Geraniaceae) were examined by light, scanning, and transmission electron microscopy. These trichomes had unicellular globular heads and stalks of different lengths and features. Two types were classified: Type I, with an elongated, large head and a short (100 μm), cylindrical stalk that was more apparent on the adaxial surface; and Type II, with a spherical, small head and a long (300μm), conical stalk that was more pronounced on the abaxial surface. The ultrastructure of secretory cells from both types was distinguished by a well-developed endoplasmic reticulum, mitochondria, plastids, dictyosomes, and numerous vacuoles that likely were involved in the storage and transport of lipophilic substances. Plasmodesmata were frequent on the walls of the secretory and stalked cells. Here, we discuss the implication of structural differentiation in these trichomes.     

Development and morphology of secretory trichomes of Calceolaria volckmanni (Scrophulariaceae)

The development and morphology of secretory trichomes of Calceolaria volckmanni was examined with light and electron microscopy. The formation and development of the glandular trichomes began with the outgrowth of a single epidermal cell which progressively increased in height and evolved into a pear-shaped cell. Subsequent divisions generated a mature trichome formed by a basal cell, a stalk "endodermal" cell and an 8-celled glandular head. Histochemical tests revealed the lipophilic nature of the secretion, the presence of terpenes and flavonoids, and displayed a particular cutinization of the walls of the stalk cell. The observed ultrastructural features of the lipophilic glandular hairs suggested the function of plastids in the secretory process.     

Glandular trichomes of <Emphasis Type="Italic">Tussilago Farfara</Emphasis> (Senecioneae,Asteraceae)

Main conclusion

The glandular trichomes are developed on the aerial organs of Tussilago farfara ; they produce phenols and terpenoids. Smooth endoplasmic reticulum and leucoplasts are the main organelles of the trichome secretory cells. The aim of this study was to characterise the morphology, anatomy, histochemistry and ultrastructure of the trichomes in Tussilago farfara as well as to identify composition of the secretory products. Structure of trichomes located on the peduncles, bracts, phyllaries, and leaves were studied by light and electron microscopy. The capitate glandular trichomes consist of a multicellular head and a biseriate long stalk. Histochemical tests and fluorescence microscopy reveal phenols and terpenoids in the head cells. During secretory stage, the head cells contain smooth and rough endoplasmic reticulum, Golgi apparatus, diversiform leucoplasts with opaque contents in lamellae, chloroplasts, mitochondria, and microbodies. In the capitate glandular trichomes of T. farfara subcuticular cavity is absent, unlike glandular trichomes in other Asteraceae species. For the first time, content of metabolites in the different vegetative and reproductive organs as well as in the isolated capitate glandular trichomes was identified by GC–MS. Forty-five compounds, including organic acids, sugars, polyols, phenolics, and terpenoids were identified. It appeared that metabolite content in the methanol extracts from peduncles, bracts and phyllaries is biochemically analogous, and similar to the metabolites from leaves, in which photosynthesis happens. At the same time, the metabolites from trichome extracts essentially differ and refer to the above-mentioned secondary substances. The study has shown that the practical value of the aerial organs of coltsfoot is provided with flavonoids produced in the capitate glandular trichomes.

     

Glandular trichomes of Ceratotheca triloba (Pedaliaceae): morphology, histochemistry and ultrastructure

This study was initiated to characterize the distribution, morphology, secretion mode, histochemistry and ultrastructure of the glandular trichomes of Ceratotheca triloba using light and electron microscopy. Its leaves bear two morphologically distinct glandular trichomes. The first type has long trichome with 8-12 basal cells of pedestal, 3-14 stalk cells, a neck cell and a head of four cells in one layer. The second type has short trichome comprising one or two basal epidermal cells, a unicellular or bicellular stalk and a multicellular head of two to eight cells. There is a marked circular area in the upper part of each head cell of the long trichome. This area is provided with micropores to exudate directly the secretory product onto the leaf surface by an eccrine pathway. The secretory product has copious amount of dark microbodies arising from plastids which are positive to Sudan tests and osmium tetroxide for unsaturated lipids. The secretion mode of short trichomes is granulocrine and involves two morphologically and histochemically distinct vesicle types: small Golgi-derived vesicles which are positive to Ruthenium Red test for mucilaginous polysaccharides; the second type is dark large microbodies similar to that of long trichomes with low quantity. These two types are stored in numerous peripheral vacuoles and discharge their contents accompanied by the formation of irregular invaginations of the plasmalemma inside the vacuoles via reverse pinocytosis. These two secretion modes of long and short trichomes are reported for the first time in the family Pedaliaceae. The long trichomes have more unsaturated lipids, while the short trichomes contain more mucilaginous polysaccharides.     

The trichomes of pericarp in Juglans (Juglandaceae): scanning microscopy, fluorescent microscopy amd histochemistry

Four types of glandular and non-glandular trichomes of pericarp in four Juglans species (J. ailanthifolia, J. cordiformis, J. mandshurica and J. regia) from Juglandaceae were studied by scanning electron microscopy, fluorescent light microscopy and histochemistry. The capitate trichomes on short stalk, the capitate trichomes on long stalk and the peltate trichomes belong to glandular trichomes; the simple hairs concern to non-glandular trichomes. The investigated species differ one from another in dimensions and distribution oftrichomes as well as the chemical content and the mechanism of secretion. The fluorescent markers and histochemical tests show the presence of flavonoids, tannins and polyphenols in trichomes on short and long stalk. In peltate trichomes the flavonoids and tannins were found in lesser quantity and the polyphenols are absent. In simple hairs the phenolic substances have not been recognized. It has been come out with the suggestion about a functional role of each type of trichomes.