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。     

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.     

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。     

Glandular Trichomes on Vegetative and Reproductive Organs of Leonotis leonurus (Lamiaceae)

The types of glandular trichomes, their ontogeny and patternof distribution on the vegetative and reproductive organs ofLeonotis leonurus at different stages of development, are studiedby light and scanning electron microscopy. Two morphologicallydistinct types of glandular trichomes (peltate and capitate)are described. Peltate trichomes, at the time of secretion,are characterized by a short stalk, which is connected witha large spherical head composed of eight cells in a single layer.Capitate trichomes can be divided into various types. Generally,they consist of a four-celled head supported by one or threestalk cells. The two kinds of trichomes differ in the secretionprocess. In the peltate trichomes, the secretory product seemsto remain accumulated in a subcuticular space, unless an externalfactor damages it. In the capitate trichomes, this product probablybecomes released through micropores. On the leaves peltate andcapitate trichomes are abundant, while on the flowers only thepeltate trichomes are numerous and the capitate are rare orabsent.Copyright 1995, 1999 Academic Press Leonotis leonurus R. Br., lion's ear, lion's tail, Lamiaceæ, glandular trichomes, morphology, ontogeny     

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.     

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.     

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.

     

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     

紫苏腺毛的形态结构和发育的研究

紫苏（Perillafrutescens（L．）Britton）叶上腺毛的研究表明：叶上腺毛主要有两种类型,一是头状腺毛,二是后状腺毛。两类腺毛都是由1个基细胞、1个柄细胞和由分泌细胞组成的头部构成。头状腺毛的头部由1个、2个或4个分泌细胞构成,其头部呈圆球形或半圆球形。盾状腺毛的头部也由1个、2个、4个或8个分泌细胞构成,其分泌细胞横向扩展使头部呈盾状。分泌盛期,大量分泌物充满角质层下间隙。两类腺毛的原始细胞均起源于叶原基或幼叶的原表皮层细胞,它通过两次平周分裂形成1个基细胞、1个柄细胞和1个头细胞,头细胞不分裂或依次进行1—3次垂周分裂,分别形成单细胞、2细胞、4细胞或8细胞的头部。     

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。     

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.     

Development and Essential Oil Content of Secretory Glands of Sage (Salvia officinalis)

Scanning electron microscopy of sage (Salvia officinalis L.) leaves confirmed the presence of two basic types of glandular trichomes consisting of a capitate stalked form containing a multicellular stalk and surmounted by a unicellular secretory head, and a capitate sessile form containing a unicellular stalk and unicellular, or multicellular, secretory head. In the latter type, secretory activity and filling of the subcuticular cavity may begin at virtually any stage of the division cycle affording fully developed glands containing from one to twelve cells in the secretory head. Gas liquid chromatographic analysis of the oil content of the most numerous gland species (capitate stalked, capitate sessile with one and with eight secretory cells) indicated only minor quantitative differences in essential oil composition. Thus, each gland type is capable of producing the four major monoterpene families (p-menthanes, pinanes, bornanes and thujanes) characteristic of sage.     

Development, oil storage and dehiscence of peltate trichomes in Thymus vulgaris (Lamiaceae)

The development, oil storage and dehiscence mechanism of the peltate oil–producing trichome in Thymus vulgaris L. was studied by conventional, fluorescence and electron scanning microscopy. A single epidermal initial forms the glandular trichome whose primordium becomes distinguishable from non–glandular trichomes at its 3–celled stage. The further divisions determine the formation of three functional compartments, (a) a basal reservoir cell, (b) an "endodermal" cell, and (c) a group of secretory head cells. At first, the gland head shows a bowl–like shape with a folded cuticular covering that raises successively assuming a dome–like form. During the differentiation of the gland cells, histochemical tests reveal the occurrence of a precise sequence of metabolic events: synthesis of RNA, proteins, glycoproteins, polysaccharides and, finally osmiophilic substances. At maturity, the cuticular sheath is formed by a non–cellulosic polysaccharide framework on which the cuticle is deposed. Proceeding towards the senescence, the polysaccharide framework disappears and a large crescent shaped pore forms, whereby essential oils are released. Collected results are discussed in order to interpret gland function and essential oil production.     

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     

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.     

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。     

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.     

甘草腺毛的形态发生和组织化学研究

利用扫描电镜及薄切片技术对甘草的腺毛形态发生和发育过程进行了观察,并对腺毛发育过程中黄酮类成分积累进行了组织化学定位研究。结果表明：甘草腺毛为多细胞构成的盾状腺毛,有长柄和短柄2种类型;前者主要分布在花萼片上,而后者主要分布于叶片上。组化鉴定结果显示：腺毛中存在着黄酮类成分、其他亲脂类和非纤维素多糖类成分;在腺毛的发育过程中,黄酮类物质是随腺毛的发育成熟,在头部盘状结构的分泌细胞及角质层下腔中积累。研究结果对进一步探讨甘草叶中黄酮类成分的合成及其作用提供科学依据。     

Cytological development and sesquiterpene lactone secretion in capitate glandular trichomes of sunflower

The secretion of sesquiterpene lactones (STL) in capitate glandular trichomes from the anther appendages of Helianthus annuus L. (Asteraceae) was observed by light and fluorescence microscopy and HPLC analysis. Disk flowers within the sunflower capitulum showed the known ontogenetic progression from the centre to the margin. During development of the florets, the trichomes in the anther appendages secreted their metabolites into the subcuticular secretion storage space in front of the two apical cells. All stages of forming the cuticular globe, from the pre-secretory to the post-secretory phase, could be observed microscopically and secretory activity was simultaneously monitored. Six germacrolides and heliangolides of known structure were selected for quantitative analysis. The increase in STL content during extension of the subcuticular space was monitored by HPLC analysis. Thereby, the start and termination of STL biosynthesis was defined in relation to other developmental stages of floret ontogenesis, particularly, the pollen formation. Part of the secreted material showed autofluorescence which could be attributed to a hydroxy-trimethoxy-flavone, as determined by NMR and mass spectroscopy. The anther trichomes were cytologically and chemically similar to foliar glandular trichomes of sunflower and represent the multicellular capitate glandular trichome type common to many Asteraceae. The ease with which anther trichomes of H. annuus can be harvested and analyzed suggests that they can provide a valuable model system for investigation of STL and flavonoid metabolism in Asteraceae.