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。     

9种榆科植物叶表皮结构特征研究

利用叶表皮离析法观察了榆科6属9种植物叶片的表皮结构。结果表明,榆科植物叶片气孔器仅分布在远轴面,不规则型,不具副卫细胞;叶片毛状体主要有腺毛和非腺毛两种类型,腺毛由基细胞、柄细胞和膨大的顶细胞构成,非腺毛均由单细胞发育而来,基部具或不具钟乳体,多数非腺毛顶部发育成长锥状,少数非腺毛顶部极短呈喙状。根据气孔器的类型和分布位置,尤其是表皮毛的基本结构和发育类型等特征,不支持将广义榆科分为两个独立科的观点。但榆科这9种植物叶表皮特征具有属间或种间差异,有一定的分类学价值。     

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。     

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.     

Comparative analysis of leaf trichome structure and composition of epicuticular flavonoids in Finnish birch species

The morphology, ultrastructure, density and distribution of trichomes on leaves of Betula pendula, B. pubescens ssp. pubescens, B. pubescens ssp. czerepanovii and B. nana were examined by means of light, scanning and transmission electron microscopy. The composition of flavonoids in ethanolic leaf surface extracts was analysed by high pressure liquid chromatography. All taxa examined contained both glandular and non-glandular trichomes (short and/or long hairs) but differed from each other in trichome ultrastructure, density and location on the leaf. Leaves of B. pubescens were more hairy than those of B. pendula, but the latter species had a higher density of glandular trichomes. Of the two subspecies of B. pubescens, leaves of ssp. pubescens had more short hairs on the leaf surface and four times the density of glandular trichomes of leaves of ssp. czerepanovii, whereas, in the latter subspecies, short hairs occurred largely on leaf veins, as in B. nana. The glandular trichomes were peltate glands, consisting of medullar and cortical cells, which differed structurally. Cortical cells possessed numerous small, poorly developed plastids and small vacuoles, whereas medullar cells had several large plastids with well-developed thylakoid systems and fewer vacuoles. In B. pubescens subspecies, vacuoles of the glandular cells contained osmiophilic deposits, which were probably phenolic, whereas in B. pendula, vacuoles of glandular trichomes were characterized by the presence of numerous myelin-like membranes. The composition of epicuticular flavonoids also differed among species. The two subspecies of B. pubescens and B. nana shared the same 12 compounds, but five of these occurred only in trace amounts in B. nana. Leaf surface extracts of B. pendula contained just six flavonoids, three of which occurred only in this species. In summary, the structure, density and distribution of leaf trichomes and the composition of epicuticular flavonoids represent good taxonomic markers for Finnish birch species.     

Trichome complement of Turnera and Piriqueta (Turneraceae)

The indumentum of Piriqueta and Turnera is made up of nine different types of trichomes, which broadly can be divided into glandular and non-glandular. Taking into account foot shape, head size and pedicle size, five variants of glandular trichomes are recognized: microcrapitate, stipitate-capitate, sessile-capitate and setiform. The non-glandular trichomes can be simple (unicellular or pluricellular-uniseriate), stellate or porrect-stellate. The setiform glandular hairs are present in most species of Piriqueta . Simple unicellular hairs are the most widespread type, frequently being found in combination with other trichomes. Stellate trichomes show a restricted distribution in both genera. Within Piriqueta , section Africana has only simple trichomes, whereas section Piriqueta has also porrect-stellate trichomes; groups of species can be set up according to the presence and type of glandular trichomes. Within Turnera the stipitate-capitate trichomes are exclusive to series Papilliferae ; sessile-capitate trichomes are found in series Microphyllae , Annulares and in some species of Salicifoliae ; clavate trichomes are found only in series Turnera ; setiform glandular hairs are exclusive to T. collotricha , whereas the microcapitate trichomes are widely distributed.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 85–97.     

Scanning electron microscopy of silica bodies and other epidermal features in Gibasis (Tradescantia) leaf

The scanning electron microscope is used to examine epidermal preparations belonging to three species of Gibasis , a genus allied to Tradescantia. The surface topography of the leaf is exposed for investigation at high magnifications, the observations add new information about structural projections from the surface and contribute to an understanding of silica deposition and its organization. Hairs of three main types occur: (1) short two-celled trichomes, hook or prickle hairs, (2) long uniseriate hairs consisting of four to six cells, (3) three-celled glandular micro-hairs. The arrangement and shape of epidermal cells and stomata are reported. Epidermal papillae were observed for the first time in two of the three species; their morphology is described in detail and their spacing expressed mathematically. Variations in the structure and distribution of silica cells are depicted, and new aspects of the silica bodies are displayed in relief. The taxonomic significance of these anatomical characters is discussed in relation to species and chromosome numbers.     

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.     

Are the trichomes in corollas of Mutisieae (Asteraceae) really twin hairs?

The corollas of three species of Onoseris , 13 species of Trichocline and one species of Uechtritzia (Asteraceae, Mutisieae) exhibit trichomes that are morphologically similar to the twin hairs which are traditionally considered exclusive of the cypselas in Asteraceae. Four types of trichomes were found: 1-celled, 2-celled, 3-celled, and 4-celled. The 3-celled and 4-celled corolla hairs are those which most resemble the typical cypsela twin hairs. The ontogeny of the corolla hairs was analysed and coincidences with the ontogeny of the cypsela twin hairs were found. (1) An anticlinal division of the epidermal mother cell originates two hair cells which, in turn, originate the basal cells (sometimes one of them is reduced or similar to the epidermal cells) by oblique or periclinal subdivision. (2) In some cases the basal cell(s) of the 3- or 4-celled corolla hair elongates and reaches the same length as the hair cells. The same kind of trichomes have been demonstrated in cypselas of Mutisieae. (3) The 1- and 2-celled corolla hairs have already been described as variants of the cypsela twin hairs. Based on this evidence we conclude that the corolla hairs of Onoseris , Trichocline and Uechtritzia are twin hairs. It is hypothesized that the 1–4 corolla hairs could be involved in water absorption, as occurs in the cypsela twin hairs.  © The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 427–433.     

两种忍冬属植物叶表皮扫描电镜观察及其生态适应性的研究

利用扫描电镜观察2种忍冬属植物的叶表皮形态特征,观察指标包括气孔器、表皮毛、表皮细胞等。扫描电镜观察结果表明:气孔仅分布在下表皮,气孔器散生并为无规则型;下表皮密被表皮毛(包括腺状毛和非腺毛);上表皮细胞的垂周壁呈沟槽状下陷;同时在下表皮还分布有瘤状的草酸钙簇晶。为进一步研究这2种忍冬属植物叶的形态解剖特征,通过光学显微镜观察发现,叶均为异面叶,栅栏组织发达,细胞长柱形;海绵组织细胞类圆形,排列紧密;叶脉机械组织发达。以上这些构造特征,都有利于减少植物体水分的散失,以适应我国西南岩溶区干旱环境特征。通过进一步分析电镜观察结果,发现在忍冬叶的上表皮中脉处有呈线状分布的腺状表皮毛,而华南忍冬叶的上表皮则无表皮毛分布,这一微形态特征可以作为二者种间特异性差别,从而为二者的合理区分提供一定的依据。     

Glandular Trichomes in Satureja thymbra Leaves

The leaves of the aromatic plant Satureja thymbra have numerousglandular trichomes of two morphologically distinct types glandularhairs and glandular scales Investigations of the anatomy ofthese glandular trichomes with serial thick sections revealedthat the glandular hairs consist of three cells a foot, stalkand head cell Glandular scales also have a unicellular footand stalk Their heads, however, are composed of 12 cells Fourof these cells are small, occupying the central region of thehead, whereas the remainder are large and peripherally arrangedMorphometric analysis showed that, in leaf surface view, glandularscales are about 17-fold larger than glandular hairs In addition,glandular scales were found to occupy 5 7 % of the entire leafsurface area In each glandular scale the total amount of essentialoil, contained within both the subcuticular space and the interiorof the secretory cells, was calculated to be 2 51 x 10^–4mm³ The volume of the essential oil produced by all glandularscales on a single mature leaf was correspondingly determinedto be 0.059 mm³ Finally, the theoretical essential oil yieldof 100 g dry leaves of S thymbra was estimated to be 3 54 %(secretory activity of glandular scales only) Satureja thymbra, glandular trichomes, morphology, morphometry     

Trichome morphology provides phylogenetically informative characters for Tremandra, Platytheca and Tetratheca (former Tremandraceae)

Summary Trichomes of Tremandra R.Br. ex DC., Platytheca Steetz and Tetratheca Sm. (Elaeocarpaceae, former Tremandraceae), together with two outgroup species of Elaeocarpus L., are illustrated using scanning electron microscopy, and their distribution on various plant organs is documented. Various trichomes types were identified that relate taxa: simple hairs, stellate hairs, short glandular trichomes, long glandular trichomes, and three forms of tubercules. Both outgroup and ingroup taxa have simple hairs. Stellate hairs are confirmed as unique to Tremandra. Prominent and sculptured multicelled tubercules, some bearing a stout hair, are characteristic of Platytheca. Smaller multicelled tubercules occur in both Platytheca and Tetratheca, except for the Western Australian taxon Te. filiformis Benth. (possibly plesiomorphic). Unicellular tubercules (papilla) characterise two species of Tetratheca. Short glandular trichomes, usually found on the ovary, also occur in both of these genera but not in all species (possibly secondary losses), while long glandular trichomes, usually on stems and leaves, occur only in some groups of Tetratheca. Within Tetratheca, Western Australian taxa that have five-merous flowers fall into three ‘groups’: seven species (together with one from South Australia) that have short glandular trichomes but no long glandular trichomes; six species that have long glandular trichomes but no short glandular trichomes; and four species that have both trichome types. All other species of Tetratheca have four-merous flowers and form two ‘groups’: 12 eastern species (including one from South Australia) that have both short glandular trichomes and long glandular trichomes; 4 western species and six eastern species that lack short glandular trichomes. On the basis of these characters, a phylogenetic hypothesis for the three genera is presented.     

Trichome morphogenesis: a cell-cycle perspective

Arabidopsis leaf hairs (trichomes) are polyploid epidermal cells with a predictable branching pattern. More than 15 genes have been identified that are involved in the regulation of branching. The cloning of the ZWICHEL, ANGUSTIFOLIA and STICHEL genes points to two mechanistic aspects of branch formation: (i) a role of the microtubule cytoskeleton; and (ii) a link to the regulation of cell divisions. The latter aspect is supported by the recent identification of an Arabidopsis mutant with multicellular trichomes, the siamese mutant, suggesting that Arabidopsis trichomes are evolutionarily derived from multicellular forms. We speculate that the spatial information for branch formation is derived from mechanisms employed in cell divisions.     

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。     

新疆委陵菜属不同花柱组植物叶表皮微形态结构的研究

利用光学显微镜和扫描电子显微镜,观察了新疆委陵菜属5组不同花柱组10种4变种植物叶表皮的微形态特征,测量统计叶表皮毛的类型、表皮细胞的形状及大小、气孔器的分布及类型、气孔的形状、大小、密度及指数、气孔外拱盖形态及其纹饰等指标。结果显示:新疆委陵菜属10种4变种植物叶的下表皮均有气孔器的分布,形状为长椭圆形、椭圆形、宽椭圆形和近圆形;气孔器的类型多为无规则四细胞型、无规则型、围绕型和辐射型;表皮毛的类型为针状毛、带状柔毛和腺毛;表皮细胞的形状分为不规则形和多边形2种类型。研究表明,新疆委陵菜属植物表皮毛特征、叶片表皮细胞的形状、垂周壁式样、气孔器的形状类型、气孔密度指数及外围蜡质纹饰等存在差异,对属以下等级的划分有重要价值,可作为物种分类及鉴别的依据,同时也为本属一些分类群间的系统关系的探讨提供佐证。     

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。     

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.     

委陵菜属锥状花柱组植物叶表皮微形态特征的研究

利用光学显微镜和扫描电子显微镜,观察了新疆委陵菜属（Potentilla L.）锥状花柱组（Sect.Conostylae(Wolf)Yü et Li）15种委陵菜植物叶表皮的微形态特征。对其叶表皮毛的类型、表皮细胞的形状及大小、气孔器的分布、气孔器类型、气孔形状、气孔大小、气孔密度、气孔指数、气孔外拱盖形态及其纹饰等指标进行分析：有几种植物叶的上表皮无气孔,而下表皮均有气孔器的分布,形状为长椭圆形、椭圆形、宽椭圆形和近圆形;气孔器的类型多为短平列四细胞型、无规则四细胞型、无规则型、围绕型和辐射型;表皮毛的类型为针状毛、带状柔毛和腺毛;表皮细胞分为不规则形和多边形;表皮毛特征、叶片表皮细胞的形状、垂周壁式样、气孔器的形状类型、气孔密度指数及蜡质纹饰等存在差异,可作为亚属间及种间分类的依据。     

Glandular hairs of Sigesbeckia jorullensis Kunth (Asteraceae): morphology,histochemistry and composition of essential oil

Long-stalked glandular hairs of outer and inner involucral bracts of Sigesbeckia jorullensis, which are important for epizoic fruit propagation, were investigated using light and scanning electron microscopy. The essential oil secreted by the hairs was analysed by chromatographic methods including gas chromatography/mass spectrometry and with a laser microprobe mass analyser. The glandular hairs consisted of a large multicellular stalk and a multicellular secreting head. The apical layer of glandular head cells was characterized by leucoplasts and calcium oxalate crystals. Below the apical cells there were up to six layers of cells containing many chloroplasts around the nucleus and surrounded by vacuoles filled with flavonoids and tannins. The essential oil originating in the head cells was secreted into the subcuticular space and may be liberated by rupture of the cuticle. It was mainly composed of sesqui- and diterpenes, with the sesquiterpene hydrocarbon germacrene-D as the main component. Monoterpenes, n-alkanes and their derivatives as well as flavonoid aglycones were also detected. The stickiness of the essential oil is probably associated with the high content of oxygenated sesqui- and diterpenes. In addition to long-stalked trichomes, small biseriate trichomes occurred, secreting small quantities of essential oil into a subcuticular space.     

黑龙江悬钩子属植物叶表皮形态结构的研究

利用扫描电子显微镜对黑龙江悬钩子属植物的叶表皮形态结构进行比较研究。结果显示:(1)悬钩子属植物叶的上表皮细胞呈多边形,垂周壁平直,或无规则形,垂周壁浅波纹;下表皮细胞无规则形,垂周壁浅波纹或深波纹。(2)表皮毛类型有单细胞直立不分支、卷曲不分支,头状腺毛和盾状腺毛四种类型。(3)气孔器均分布于下表皮,且气孔器类型为无规则形;气孔外拱盖单层、内缘平滑或不规则波状。研究表明,黑龙江悬钩子属植物的叶表皮微形态学特征表现出一定差异性,对种间的划分和鉴定具有一定的分类学意义。