Morphology, ontogeny and histochemistry of secretory trichomes of Geranium robertianum (Geraniaceae)

Geranium robertianum bears three types of glandular uniseriate trichomes which originate from a single protodermal cell and develop through periclinal divisions. Type I trichomes are procumbent and have an oval apical cell, two stalk cells and a basal cell. Type II trichomes are erect and have a pear shaped apical cell, two stalk cells and a basal cell. Type III trichomes are much longer than the other two types and have an elongated apical cell, five long stalk cells and a basal cell. Type I and type II trichomes are common on leaves while III trichomes are more abundant on flower structures.
Type I and type II trichomes secrete terpenoids and phenols. Type III trichomes are characterized by the accumulation of anthocyanins in the apical cell and secrete flavonoids.     

A new species of Ocimum (Lamiaceae) from Swaziland

A new species of Ocimum , O. motjaneanum McCallum & K. Balkwill, endemic to serpentine soils in the north-west of Swaziland, is described and compared with O. obovatum, the species it most closely resembles. It has a very localized distribution with an area of occupancy under threat of encroachment by exotic trees and development, which makes it critically endangered (CR B1a + b(iii)) using the IUCN guidelines. We recommend habitat protection and incorporation of one site into an existing nature reserve to assure the continuation of this species.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 379–383.     

Glandular trichomes and essential oil constituents ofCalamintha menthifolia (Lamiaceae)

Plant material ofCalamintha menthifolia was collected from two populations grown in the Vikos-Aoos area (NW. Greece). The structure and ontogeny of the glandular trichomes and the chemical composition of the secreted essential oil were studied. The leaves ofC. menthifolia have numerous glandular trichomes of three morphological distinct types: 1) glandular scales with a unicellular foot and stalk, and a 12-celled head, 2) glandular hairs with a unicellular foot and stalk, and a unicellular pear-shaped head and 3) glandular hairs with a unicellular foot, a bicellular stalk and a unicellular elongated head. Qualitative and quantitative GC-MS analyses of the essential oils revealed piperitone oxide as the main constituent in both populations. In the light of these results the relationships beweenC. menthifolia and other members of theSatureja group as well as its taxonomy are discussed.     

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.     

Development,structure, and occurrence of secretory trichomes ofPharbitis

Summary Secretory trichomes develop from epidermal cells on the leaf primordia and stem ofPharbitis nil. Following an initial growth phase, trichomes begin active secretion of a protein-carbohydrate mucilage. This mucilage covers the shoot apex and developing leaves ofPharbitis.The secretory cells possess cellular organelles in forms usually associated with actively secreting cells: many mitochondria, an elaborate network of rough endoplasmic reticulum (RER), many free ribosomes, and numerous dictyosomes. The role of the dictyosomes is twofold: 1. dictyosome vesicles bud coated vesicles which transport materials from the cell and, 2. dictyosome vesicles coalesce, forming large storage vesicles. The storage vesicles are surrounded by, and often in contact with, poculiform RER. The RER forms an interconnected network throughout the cytoplasm, extending from the nuclear envelope to the plasmalemma. Distended profiles of RER are frequently in direct contact with the plasmalemma. Thus, inPharbitis secretory trichomes, it is the coated vesicles and RER which are active in secretion export. These findings imply a secretory pathway which deviates from the usual pattern in glandular cells.Predoctoral fellow of National Science Foundation during part of the investigation.     

Morphology and histochemistry of the glandular trichomes of Lippia scaberrima (Verbenaceae)

BACKGROUND AND AIMS: Lippia scaberrima, an aromatic indigenous South African plant, with medicinal application, potentially has economic value. The production of essential oil from this plant has not been optimized, and this study of the chemico-morphological characteristics was aimed at determining the location of oil production within the plant. Furthermore, the locality of other secondary metabolites important in medicinal applications needed to be ascertained. This information would be useful in deciding the protocol required for isolation of such compounds. METHODS: The morphology of the glandular trichomes was investigated using a combination of scanning electron and light microscopy. Concurrently, the chemical content was studied by applying various chemical reagents and fluorescence microscopy. KEY RESULTS: Three types of trichomes were distinguished on the material investigated. Large, bulbous peltate glands containing compounds of terpenoid nature are probably the main site of essential oil accumulation. Small glands were found to be both peltate and capitate and fluorescent stain indicated the possible presence of phenolic compounds. The third type was a slender tapered seta with an ornamented surface and uniseriate base, and evidently secretory in nature. CONCLUSIONS: This study linking the chemical content and morphology of the glandular trichomes of L. scaberrima has contributed to the knowledge and understanding of secretory structures of Lippia spp. in general.     

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.     

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.     

Effects of light intensity on the anatomical structure,secretory structures,histochemistry and essential oil composition of Aeollanthus suaveolens Mart. ex Spreng. (Lamiaceae)

Aeollanthus suaveolens Mart. ex Spreng belongs to Lamiaceae family, and in the Amazon this species is cultivated by natives’ people, this medicinal plant is popularly known as Catinga-de-mulata, being used by the population for general pain treatment. The present study analyzed the effects of light intensity on the anatomy, secretory structures, histochemistry and composition of essential oil of leaf and stem of A. suaveolens. The anatomical structure were observed in response to two light intensities, namely 50% (half shade, HS) and 100% (full sun, FS) light. Histochemical analyses were performed to detect lipids, essential oils, alkaloids, phenolic compounds, sesquiterpene lactones, mucilage, and tannins. Secretory structures were observed under a scanning electron microscope. The results obtained in the present work indicate that the light intensity can affect the histochemistry and structures of A. suaveolens. Cross sections of the leaves and stem revealed glandular trichomes on both leaf surfaces as well as the stem surface. Essential oil was detected by histochemical analyses in all types of secretory trichomes. These anatomical and histochemical responses suggest modifications to protect the photosynthetic apparatus from excess light, in addition we note that in the chemical composition of the essential oil the class of hydrocarbons sesquiterpene prevailed.     

Diversity,phytochemical and medicinal potential of the genus Ocimum L. (Lamiaceae)

Phytochemistry Reviews - Ocimum, commonly known as Tulsi, is a huge genus within family Lamiaceae, comprising about 64 species of annual to perennial aromatic medicinal herbs with a long history of...     

Quantitative taxonomic analyses of Actinidia (Actinidiaceae) in China based on micromorphological characters of foliar trichomes

There are various arguments on classification of the genus Actinidia Lindl., a genus with approximately 63 species, 59 of which have been found in China. The paper investigated the characteristics of foliar trichomes of 35 taxa from China under optical microscope, including size, celluar structure, distribution and density. According to their micromorphological characteristics, foliar trichomes can be classified into the following six categories: 1) single-cell hairs; 2) uniseriate hairs, including linear, bulbous, twisted, straight-walled, and bent-walled hairs; 3) multiseriate hairs, including twisted, straight-walled and gradually sharpening, straight-walled and suddenly sharpening, bent-walled and gradually sharpening, and suddenly sharpening hairs; 4) multiseriate thick hairs, including pillar hairs, gradually sharpening thick hairs, and suddenly sharpening thick hairs; 5) stellate hairs, including parenchyma-stellate and sclerenchyma-stellate (normal state and special states such as rosulate, peltate-stellate, and overlopping-stellate) hairs; and 6) dichotomous hairs. On the basis of the micromorphological characteristics of foliar trichomes in Actinidia, with Clematoclethra lasioclada as an outgroup, both the quantitative cladistic analysis and phenetic analysis were performed using Wagner method and UPGMA clustering method respectively to reconstruct the phylogeny of Actinidia in China. The phylogenetic tree generated by cladistic analysis suggests that the sect. Leiocarpae be a monophyletic group, but other three sections, i.e., sect. Maculatae, sect. Strigosae and sect. Stellatae, be non-monophyletic groups. The results obtained from the phenetic analysis reflect relationships among the taxa of Actinidia in China, especially a close relationship between A. chinensis and A. deliciosa, and a relatively remote relationship between A. callosa var. henryi and A. callosa var. discolor. In conclusion, the micromorphological characters of foliar trichomes and the methods of quantitative taxonomic analysis are of key importance tostudies on phylogenetic and phenetic relationships of Actinidia.     

The anatomical and micromorphological properties of three endemic and medicinal Salvia species (Lamiaceae) in Erzincan (Turkey)

In this study, we comparatively investigated three endemic Salvia species spreading in Erzincan (Turkey) in terms of anatomy and micromorphology. For anatomical investigation, cross sections taken from stems and leaves of the species were examined under light microscope. For micromorphological investigation, epidermal surface and nutlet structure were examined using scanning electron microscopy. The S. euphratica and S. divaricata species were examined anatomically and micromorphologically for the first time, and S. hypargea was examined micromorphologically for the first time. In anatomical examinations, it was seen that stem and leaf structures of the species were similar. In micromorphological analyses, it has been seen that hairiness of the nutlet surface and nutlet ornamentations (verrucate and rugose type) created a difference between the species.     

The essential oil secretory structures of Prostanthera ovalifolia (Lamiaceae)

The structure of the essential oil secretory tissues of Prostanthera ovalifolia R.Br was investigated using bright- and dark-field optical microscopy, and scanning and transmission electron microscopy. The leaves of P. ovalifolia have glandular trichomes of the peltate type common to many Lamiaceae species. The trichomes consist of a basal cell embedded in the epidermis, a stalk cell with heavily cutinized walls and a 16-celled secretory head, but they differ from those of many previously reported Lamiaceae species in their morphological form defined by the elevated cuticle. The sub-cuticular space contains a mixture of lipid and aqueous phases. Secretory cells have dense cytoplasm with many leucoplasts present. Volatile terpenoids are eliminated from the cytoplasm into the sub-cuticular space, the site of essential oil accumulation, via granulocrine secretion.     

Evolution of trichomes and its systematic significance in Salvia (Mentheae; Nepetoideae; Lamiaceae)

We have investigated the trichome characteristics in representative species of Salvia and Pleudia in order to evaluate this source of morphological evidence for addressing problems regarding generic delimitation and subgeneric classification. Trichomes of 46 Salvia spp., representing three subgenera in Iran, were investigated using light microscopy and scanning electron microscopy. General trichome characteristics were constant among different populations of a certain species, but showed a degree of variability useful in the delimitation of taxa, specifically at lower taxonomic levels. Trichome characters of taxonomic interest are as follows: types of glandular hair; number of composing cells (uni‐, bi‐ or multicellular); size and thickness; branching pattern; and presence of papillae on the surface. Non‐glandular trichomes can be simple and branched. Glandular trichomes can be stalked, subsessile or sessile. Our investigation reveals the usefulness of such characters in providing fundamental taxonomic criteria for taxon delimitation in these genera at various levels, especially at the specific rank. Furthermore, the data presented here indicate the potential applicability of such characters in the determination of evolutionary trends in Salvia and allies. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 180 , 241–257.     

中国猕猴桃属植物叶表皮毛策形态特征及数量分类分析

选取国产猕猴桃３５个分类群的代表植株,应用光学显微对其新鲜叶表皮毛的微形态特征,形体大小、细胞结构、分布和密度等多态性状和数量性状进行了观察和测量,该属植物的叶表皮毛微形态特征可归纳为６个类型：１）单细胞毛;２）单列多细胞毛,每形单列毛、泡状单列毛、扭曲毛、直壁单列毛、曲壁单列毛;３）多列渐尖毛和急尖毛,包括多列曲壁渐尖毛和急尖毛,多列直壁渐尖毛和急尖毛;４）多列粗毛,包括柱状毛,多列渐尖粗毛,多     

Glandular Hairs and Essential Oil in Developing Leaves of Ocimum basilicum L. (Lamiaceae)

The types of glandular hairs and their pattern of distributionon leaves of Ocimum basilicum L. at different stages, were investigated.Their essential oil at the various stages was chemically analysed.The density of the glandular hairs appears to be very high onyoung meristematic leaves and on meristematic regions of olderleaves. When the leaf regions start to expand no new glandularhairs are produced and their density decreases. The essentialoil differs in the percentage of some of its components betweenyoung and mature leaf regions.Copyright 1993, 1999 AcademicPress Ocimum basilicum, sweet basil, glandular hairs, leaf development, essential oil, cugenol, methyl chavicol, 1,8-cineole     

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.     

Composition of the essential oils of Ocimum urticifolium (Lamiaceae) chemotypes grown in Rwanda

NTEZURUBANZA, L., SCHEFFER, J. J. C. & SVENDSEN, A. B., 1988. Composition of the essential oils of Ocimum urticifolium (Lamiaceae) chemotypes grown in Rwanda. Leaves and flowers of 48 specimen of Ocimum urticifolium (synonym: O. suave), Lamiaceae, were collected in different parts of Rwanda. The volatile components of each specimen were isolated by hydrodistillation under standardized conditions, and the 48 essential oils analysed by thin-layer chromatography. This preliminary analysis showed six different oil compositions. Thereafter, six oil samples were fractionated by liquid—solid chromatography. The fractions thus obtained, and all other oil samples were then analyzed by capillary gas chromatography. In this way 46 compounds were identified in the various oil samples. Eight components, which amounted to more than 10% in at least one oil sample, were used to characterize the oils. Twelve oil types were distinguished, three of which were dominated by one component only: eugenol, methyleugenol and transmethylisoeugenol respectively. The other types were more complex. In some parts of Rwanda almost one oil type seemed to occur, whereas in other areas a more heterogeneous collection of types was found. Eugenol was quantitatively most important in 31 of the 48 oil samples, in 22 of which this component amounted to more than 50%.     

Pericarp structure and phylogeny of tribe Mentheae (Lamiaceae)

Pericarp structure is investigated in 143 specimens belonging to 59 out of ca. 65 genera and 134 out of ca. 2,000 species of Lamiaceae subfamily Nepetoideae tribe Mentheae. Some of the data presented by earlier authors are believed to be incorrect. The variation in pericarp structure has been found to be strongly correlated to the variations in DNA and gross morphological characters. The groups of two-staminate genera (Salvia, etc.) and four-staminate genera of Salviinae (Lepechinia and Chaunostoma) can be distinguished from each other and from other labiates by differences in pericarp structure. The other two subtribes, Menthinae and Nepetinae, differ considerably, but their variations strongly overlap. The differences in pericarp structure also suggest that Menthinae can be divided into three main monophyletic groups based on other data: (1) the main bulk of the tribe, (2) a group of Prunella and Cleonia and perhaps also Horminum, and (3) the characteristic genus Lycopus. Pericarp data disagree with a suggestion that the genus Melissa should be included in the subtribe Salviinae. Modifications of the subtribal classification of Mentheae are discussed.