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。     

Foliar secretory trichomes of Ocimum obovatum (Lamiaceae): micromorphological structure and histochemistry

This study characterises the micromorphology, ultrastructure and main chemical constituents of the foliar glandular trichomes of Ocimum obovatum using light and electron microscopy and a variety of histochemical tests. Two types of glandular trichomes occur on the leaves: large peltate and small capitate. The head of each peltate trichome is made up of four broad head cells in one layer. The head of each capitate trichome is composed of two broad head cells in one layer (type I) or a single oval head cell (type II, rare). In peltate heads, secretory materials are gradually transported to the subcuticular space via fracture in the four sutures at the connecting walls of the head cells. Release to the head periphery occurs through opposite fracture in the four sutures in the head cuticle. In type I capitate trichomes, release of the secretions to the subcuticular space occurs via a pore between the two head cells, and release to the head periphery occurs through the opposite pore in the head cuticle. In type II capitate trichomes, the secreted material is released from the head cell through a ruptured particular squared area at the central part of the head cuticle. These secretion modes are reported for the first time in the family Lamiaceae. Histochemical tests showed that the secretory materials in the glandular trichomes are mainly essential oils, lipophilic substances and polysaccharides. Large peltate trichomes contain a large quantity of these substances than the small capitate trichomes. Ultrastructural evidence suggests that the plastids produce numerous lipid droplets, and the numerous polysaccharide small vesicles are derived from Golgi bodies.     

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。     

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。     

Structural and Histochemical Investigation of the Glandular Trichomes of Salvia aurea L. Leaves, and Chemical Analysis of the Essential Oil

Anatomical and histological investigations of the secretoryhairs ofSalvia aurea leaves, and identification of the maincomponents of the essential oil were carried out. Two typesof glandular trichome were found: peltate glands, characterizedby a short stalk and a large six to eight-celled head, and capitatetrichomes which were further subdivided into two kinds, thefirst with a short monocellular stalk and two-cellular head(type I), and the second with a multicellular stalk, a neckcell and a small globose unicellular head (type II). Whereaspeltate glands and type I capitate trichomes were always present,type II capitate glands were not found in all leaf samples.The histochemical study suggested an ‘endodermal’role for the stalk cell (peltate and capitate type I) as wellas for the neck cell (capitate type II), preventing the lossof essential oil. Histological reactions also revealed the complexnature of the material secreted by all types ofS. aurea trichome,including polysaccharides, polyphenols and proteins, in additionto the essential oil. Qualitative and quantitative GC-MS analysisof the essential oil revealed camphor to be the main constituent.The findings are discussed in relation to studies of trichomesfrom other members of the Lamiaceae. Salvia aurea L.; glandular trichomes; histochemistry; essential oil     

Glandular Trichomes on the Leaves and Flowers of Plectranthus ornatus: Morphology, Distribution and Histochemistry

The types of glandular trichomes and their distribution on leavesand flowers of Plectranthus ornatus were investigated at differentstages of their development. Five morphological types of glandulartrichomes are described. Peltate trichomes, confined to theleaf abaxial surface, have, in vivo, an uncommon but characteristicorange to brownish colour. Capitate trichomes, uniformly distributedon both leaf surfaces, are divided into two types accordingto their structure and secretory processes. In long-stalkedcapitate trichomes, a heterogeneous secretion (a gumresin) isstored temporarily in a large subcuticular space, being releasedby cuticle rupture, whereas, in the short-stalked capitate trichomes,the secretion, mainly polysaccharidic, is probably exuded viamicropores. On the leaves, digitiform trichomes, which do notshow a clear distinction between the apical glandular cell andthe subsidiary cells, occur with a similar distribution to thecapitate trichomes. They do not develop a subcuticular space,and secrete small amounts of essential oils in association withpolysaccharides. The reproductive organs, particularly the calyxand corolla, exhibit, in addition to the reported trichomes,unusual conoidal trichomes with long unicellular conical heads.A large apical pore, formed by tip disruption, releases thesecretion (a gumresin) stored in a rostrum-like projection.On the stamens and carpels, digitiform, capitate and conoidaltrichomes are absent, but peltate trichomes are numerous. Theyoccur between the two anther lobes, on the basal portion ofthe style, and between the four lobes of the ovary. The resultspresented are compared with those of other studies on Lamiaceaeglandular trichomes. Copyright 1999 Annals of Botany Company Plectranthus ornatus Codd, Lamiaceae, glandular trichomes, morphology, histochemistry, essential oils and mucilage secretion.     

Trichome micromorphology in Turkish species of Ziziphora (Lamiaceae)

Ziziphora L. is represented by 5 species and 2 subspecies in the flora of Turkey: Z. clinopodioides, Z. capitata, Z. persica, Z. tenuior, Z. taurica subsp. taurica, Z. taurica subsp. cleonioides. It is difficult to distinguish between some Ziziphora taxa because of their morphological similarities. In this study, the leaf and calyx trichomes of Ziziphora taxa in Turkey were studied in order to assess anatomical variations that may serve as distinguishing characters. Their micromorphological features were surveyed by scanning electron microscopy (SEM) and light microscopy (LM). Trichomes on leaves and calyx can be divided into two general types: non‐glandular trichomes and glandular (secretory) trichomes. The non‐ glandular trichomes are simple, acicular or curved with cuticular micropapillae. They usually consist of one or more additional cells. The glandular trichomes are divided into two types: peltate and capitate and Ziziphora taxa can easily be distinguished by presence/absence, density and types of glandular trichomes on leaves and calyx. The peltate trichomes consist of 12 or 18 secretory head cells in a single disc; four or six central cells surrounded by eight or twelve peripheral ones. Peltate trichomes are absent on the adaxial leaf surface of Z. capitata and Z. persica. Two types of capitate trichomes are present in Ziziphora. The capitate trichomes are only absent on the calyx surface of Z. persica. In addition, the trichome micromorphology provides some support for separating the two subspecies of Z. taurica. In conclusion, Ziziphora taxa can easily be distinguished by cell number, cell shape presence/absence and density of the glandular trichomes on leaves and calyx.     

Localization of salvinorin A and related compounds in glandular trichomes of the psychoactive sage, Salvia divinorum

BACKGROUND AND AIMS: Salvia divinorum produces several closely related neoclerodane diterpenes. The most abundant of these, salvinorin A, is responsible for the psychoactive properties of the plant. To determine where these compounds occur in the plant, various organs, tissues and glandular secretions were chemically analysed. A microscopic survey of the S. divinorum plant was performed to examine the various types of trichomes present and to determine their distribution. METHODS: Chemical analyses were performed using thin layer chromatographic and histochemical techniques. Trichomes were examined using conventional light microscopy and scanning electron microscopy. KEY RESULTS: It was found that neoclerodane diterpenes are secreted as components of a resin that accumulates in peltate glandular trichomes, specifically in the subcuticular space that exists between the trichome head cells and the cuticle that encloses them. Four main types of trichomes were observed: peltate glandular trichomes, short-stalked capitate glandular trichomes, long-stalked capitate glandular trichomes and non-glandular trichomes. Their morphology and distribution is described. Peltate glandular trichomes were only found on the abaxial surfaces of the leaves, stems, rachises, bracts, pedicles and calyces. This was consistent with chemical analyses, which showed the presence of neoclerodane diterpenes in these organs, but not in parts of the plant where peltate glandular trichomes are absent. CONCLUSIONS: Salvinorin A and related compounds are secreted as components of a complex resin that accumulates in the subcuticular space of peltate glandular trichomes.     

Glandular Hairs of Salvia officinalis: New Data on Morphology, Localization and Histochemistry in Relation to Function

The structure, site and histochemistry of glandular hairs onthe vegetative and reproductive parts ofSalvia officinalis wereinvestigated by UV and conventional light microscopy and byscanning electron microscopy. Five distinct types of glandularhair (one peltate and four capitate) with different sites, secretorymodes and secretions, were identified, and a functional rolepostulated for each type. All the hair types show mixed secretions,i.e., hydrophilic and lipophilic, except type I capitate hairs,which have hydrophilic secretions only. In peltate hairs andin type II capitate hairs hydrophilic secretion prevails; inthe remaining types, lipophilic secretion dominates. The manner,time and role of erection of peltate hairs on the reproductiveorgans and the role of non-glandular hairs are also considered.Copyright 1999 Annals of Botany Company Salvia officinalis, Labiatae, rising of peltate hairs, capitate hairs, secretion, plant/insect interaction.     

Sideritis syriaca ssp. syriaca: Glandular trichome structure and development in relation to systematics

Sideritis syriaca ssp. syriaca is a taxon with a low essential oil content. Its leaves bear glandular trichomes of two types: long hairs with a 4-celled head, a 4-celled stalk and a 4-celled foot (reported for the first time in Lamiaceae) and short hairs with a 4–celled head, a unicellular stalk and a unicellular foot. The second type is considered intermediate between the capitate and peltate hairs, common in Lamiaceae, but found in S. syriaca ssp. syriaca. The ontogeny of the trichome types is described. The possible significance of the glandular trichome structure to Lamiaceae systematics is further discussed.     

羽叶薰衣草表皮毛的发育解剖学研究

对羽叶薰衣草(LavandulapinnataL.)茎和叶上两种表皮毛(腺毛和非腺毛)发育的解剖学观察表明,两者的发生都源于茎或叶的原表皮细胞,但外部形态、发育过程及功能明显不同。腺毛有头状腺毛和盾状腺毛两种类型,均由1个基细胞、1个柄细胞和头部细胞构成。头状腺毛的头部只有1个或2个分泌细胞,盾状腺毛由8个分泌细胞构成头部。非腺毛由3-20个细胞组成,可分为三种类型:单列不分枝、二叉分枝和三叉及三叉以上多分枝的树状分枝。非腺毛的顶部细胞由基部到顶部逐渐变细,先端成尖形。腺毛发育由原表皮细胞经两次平周分裂形成,由于柄细胞和头部细胞所处的分化状态不同而发育成两类腺毛。非腺毛由非腺毛原始细胞经二次或多次平周分裂和不均等分裂,再发育成数个至二十多个子细胞。     

Glandular trichomes of Humulus lupulus var. Brewer's Gold: Ontogeny and histochemical characterization of the secretion

Leaves of Humulus lupulus possess two types of glandular trichomes: - peltate (lupulin) and bulbous.
Peltate trichomes are formed from a protodermal cell by two anticlinal divisions in perpendicular planes, followed by two periclinal ones that give rise to the initials of the head cells, the basal and the stalk cells. Head cells divide successively in radial and irregular planes. Fully developed peltate trichomes are built of a glandular head consisting of 30 to 72 cells, four stalk cells and four basal cells.
Bulbous trichomes are also formed from a protodermal cell by an anticlinal division followed by two periclinal ones that produce the initials of the glandular head cells, and the basal and stalk cells. Fully developed bulbous trichomes consist of four (occasionally eight) head glandular cells, two stalk cells and two basal cells.
The density of peltate trichomes decreases with the expansion of the leaves.
Both peltate and bulbous trichomes secrete essential oils. Peltate trichomes are the preferential site for the synthesis of bitter resins. Tannic acids could not be detected histochemically either in peltate or in bulbous trichomes. Both types of trichomes produce secretion that accumulates in the subcuticular space, being released, in the case of bulbous trichomes, by rupture of the cuticle.     

Developmental Ultrastructure of Glandular Trichomes of <Emphasis Type="Italic">Rosmarinus officinalis</Emphasis>: Secretory Cavity and Secretory Vesicle Formation

Glandular trichomes in the leaf lamina of Rosmarinus officinalis L. were examined by scanning and transmission electron microscopy. The leaves were characterized by an abundance of two types of glandular trichomes—small capitate and large peltate glandular trichomes. In addition to the glandular trichomes, numerous non-glandular trichomes were present on the abaxial surface of the leaf. These trichomes mainly predominated on the midrib, whereas glandular trichomes occurred on non-vein areas. At the initial phase of secretory cavity formation, hyaline areas were abundant in periclinal walls of head cells, while they were not observed in the anticlinal walls. The hyaline areas gradually increased in size, fusing with other areas throughout the wall. Loose wall material adjacent to hyaline areas was released from the head cell walls and migrated into the secretory cavities. As the secretory cavities continued to enlarge, the new vesicles emerging into the secretory cavities from the walls of head cells became surrounded with the surface of a typical membrane. They developed a round shape, but the contours of the vesicle surfaces appeared polygonal when tightly packed inside a cavity. These vesicles varied in size; small vesicles often possessed electron-dense contents, while large vesicles contained electron-light contents.     

An investigation of the foliar trichomes of Tetradenia riparia (Hochst.) Codd [Lamiaceae]: An important medicinal plant of Southern Africa

The morphology and distribution of leaf trichomes of Tetradenia riparia were studied using light and scanning microscopy. Three morphologically distinct types of trichomes were observed on T. riparia leaf surfaces: glandular capitate (short and long stalked), peltate and non-glandular. The glandular and non-glandular trichomes were present in abundance on both the adaxial and abaxial surfaces. Young leaves were densely covered with trichomes; however, the density of trichomes decreases progressively with leaf maturity. This suggests that the trichomes are established early in leaf differentiation and their density decreases with leaf development and age.     

兰花鼠尾草(Salvia farinacea Benth.)两类腺毛发育过程中超微结构的研究

电镜观察表明：兰花鼠尾草（Salvia farinacea Benth.）的头状腺毛（capitate trichomes）和盾状腺毛(peltate trichomes）在超微结构方面存在明显不同,盾状腺毛的分泌细胞中占优势的细胞器是质体,而头状腺毛中是内质网和质体;成熟的盾状腺毛角质层下间隙明显,而头状腺毛不明显;盾状腺毛的柄细胞的侧壁出现完全的角质化现象,而头状腺毛则无;头状腺毛的基细胞液化程度比盾状腺毛的高。     

Structural investigation of the glandular trichomes of <Emphasis Type="Italic">Salvia argentea</Emphasis>

The morphology, anatomy and distribution of glandular trichomes on the aerial organs of Salvia argentea L. has been investigated. Two morphologically distinct types of glandular trichomes were determined. Capitate glandular trichomes forming a base 1–7 celled, a stalk 1–5 celled or no stalk and a head uni- or bicellular had various types. In capitate trichomes, the neck cell that has an important role especially for xeroformic plants, acting to prevent the backflow of secreted substance through the apoplast has been distinctively observed in the investigated species. The capitate trichomes were present abundantly on all aerial organs of S. argentea. Peltate glandular trichomes had a large secretory head forming 1–5, 8 central and 8–10, 12, 14 peripheral cells. Peltate trichomes are present on all aerial organs, except petiole, being the most abundant on calyx and corolla. Results were shown by tables and photographs.     

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.     

紫苏腺毛的形态发生研究

紫苏叶上有两种腺毛:盾状腺毛和头状腺毛。两者都具1个基细胞、1个柄细胞和头部。前者的头部可由1、2、4或8个分泌细胞组成,扩展成盾状;后者的头部由1、2或4个分泌细胞组成,聚成圆球状。两种腺毛的原始细胞都来源于原表皮细胞,经两次平周分裂产生基细胞、柄细胞和顶细胞。在腺毛后期的形态发生中,柄细胞的分化状态决定腺毛的类型。若柄细胞保持扁平状且处于分生状态时,其顶细胞将发育成盾状腺毛的头部;若柄细胞纵向引长并迅速液泡化时,其顶细胞将发育成头状腺毛的头部。     

紫苏腺毛的形态发生研究

紫苏叶上有两种腺毛：质状腺毛和头状腺毛。两者都具１个基细胞、１个柄细胞和头部。前者的头部可由１、２、４或８个分泌细胞组成,扩展成质状;后者的头部由１、２或４个分泌细胞组成,聚成圆球状。两种腺毛的原始细胞都来源于原表皮细胞,经两次平周分裂产生基细胞、柄细胞和顶细胞。在腺毛后期的形态发生中,柄细胞的分化状态决定腺毛的类型。若柄细胞保持扁平关且处于分生状态时,其顶细胞将发育成质状腺毛的头部;若柄细胞纵向     

Insight into the structure and chemistry of glandular trichomes of Labiatae, with emphasis on subfamily Lamioideae

Glandular trichomes of Labiatae are among the most investigated secretory structures. Most species studied belong to subfamily Nepetoidae, including plants with aromatic properties, while so far a few species of subfamily Lamioideae were examined. In this work, we studied the micromorphology, ultrastructure, type and release of secretion of the glandular trichomes present on leaves and flowers of several species belonging to subfamily Lamioideae, (Stachys alopecuros (L.) Bentham subsp. alopecuros, S. officinalis (L.) Trevisan subsp. officinalis, S. germanica L. subsp. germanica, S. germanica L. subsp. salviifolia (Ten.) Gams, S. sylvatica L., S. heraclea All., S. plumosa Griseb., S. annua L., Prasium majus L., Sideritis romana L.) and one to the sister group Scutellarioideae (Scutellaria galericulata L.). Besides the well-known peltate and small capitate trichomes, widely described in the literature, other types of glandular trichomes were encountered; stalked peltate hairs and large capitate hairs. In particular, a new type of capitate trichome, exclusive of calices and corollas, which presents a mode and release of secretion never described before, is reported.