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.     

Cell Expansion and Wall pH in the Fern Regnellidium diphyllum, A Plant Lacking Acidinduced Growth

Petioles of the semi-aquatic fern Regnellidium diphyllum donot show acid growth but low wall pH is a necessary conditionfor maximum rates of IAA-induced cell expansion. Measurementsof wall pH by two indirect methods indicate an unusually lowvalue, in the range pH 4 to 5. This is one to two pH units belowthat estimated for petioles of the semi-aquatic dicotyledonNymphoides peltata, a species in which IAA aand ethylene causegrowth responses very similar to those in Regnellidium but whereacid growth occurs. Having shown previously that fusicoccinenhances proton secretion in both Regnellidium and Nymphoides,we now show that although it causes a reduction in the estimatedapoplast pH to below 4·0 in Regnellidium, cell expansionis not promoted. The FC-induced reduction in pH in Nymphoidesis less and occurs more slowly, but growth is promoted significantly;when IAA and fusicoccin are present together, growth promotionis approximately additive for Nymphoides A model is proposed for Regnellidium in which equilibrium wallpH is maintained at a low value that is optimal for acid growth,the availability of acid-labile sites in the wall being thechief limitation to cell extension. We suggest that this controlmechanism may be widespread for organs without a cuticle, includingroots and the gametophytes of lower plants growing in acidicconditions. Key words: Acid growth, wall pH, fusicoccin, Regnellidium diphyllum, Nymphoides peltata     

Calcium-Dependent Lamina Production of Nymphoides peltata (Gmel.) O. Kuntze (Menyanthaceae): Implications for Distribution

Nymphoides peltata (Gmel.) O. Kuntze, a nymphaeid macrophyte,occurs commonly in polder and fluviatile areas in large partsof Europe and Asia. In contrast to the nymphaeid macrophytesNymphaea alba L. and Nuphar lutea (L.) Sm., Nymphoides peltatais almost completely absent from poorly-buffered waters andis never found in acid water bodies. Transplantation experimentsin water bodies of varying alkalinity demonstrated that, irrespectiveof the sediment type, leaf production of Nymphoides did occurin poorly-buffered waters, but not in acid waters. Cultivation experiments showed that floating leaf developmentof Nymphoides peltata could only take place if sufficient calciumwas available in the water layer or in twice-demineralized water.Addition of calcium to an acid cultivation medium or to watercollected from an acid moorland pool resulted in leaf production.Growth of Nymphoides in acid waters is impossible due to insufficientcalcium concentrations in the water layer of such waters. Itis suggested that the absence of Nymphoides peltata in somepoorly-buffered water bodies is partly due to the spatial isolationfrom rivers and canals and the high frequence of desiccation.The restricted occurrence of Nymphoides peltata to well-bufferedalkaline waters is functionally more related to the calciumavailability than to the bicarbonate content. Key words: Aquatic macrophytes, distribution, Nymphoides peltata, leaf production, calcium, acid, poorly-buffered and alkaline water     

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.     

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.     

The Ultrastructure of the Glandular Trichomes of Solanum tuberosum

The potato plant has two types of glandular trichomes whichwere investigated by electron microscopy. One type has a eight celled globular head on a neck cell anda stalk cell Each glandular cell has many rather large vacuoles,a large nucleus, many ribosomes and mitochondria, a few Golgibodies, and darkly coloured, often irregular plastids (chloroplasts).The plastids are mostly located near the axial cell wall borderinga large central intercellular space filled with secretion materialThe plastids are assumed to participate in the formation ofthe secretion material, which reacts positively to esterasetests. The outer wall is covered by a thin cuticle. The other type has a club-shaped multicellular head on a singlestalk cell. The cytoplasmic features in the cells are similarto those of the globular-headed trichome, except that they possesslarge central vacuoles and randomly distributed plastids. Centricendoplasmic reticulum has been observed in young cells. Intercellularspaces develop between the cells and into the outer wall, whichis thus split into two. Whereas the older glandular cells reactpositively to tests for esterase, the secretion material itselfis pectinaceous and reacts negatively. The outer wall is cutinizedand covered by a cuticle. Solanum tuberosum L., potato, glandular trichomes, ultrastructure     

The Ligule of Isoetes lacustris: Ultrastructure, Mucilage Composition, and a Possible Pathway of Secretion

Mature ligules of Isoetes lacustris can be divided anatomicallyinto three ultrastructurally different regions. First, the basalregion contains large numbers of two types of protein bodies.Second, the cells of the sub-marginal region are characterizedby a well developed Golgi apparatus closely associated withthe tubular compartments of the endoplasmic reticulum. Third,the peripheral region consists of one to three layers of cellsshowing an extremely well developed rough asternal and smoothtubular endoplasmic reticulum (ER). The tubular ER compartmentsare frequently observed in close attachment to the plasmalemma.The outermost peripheral cells are covered with a mucilaginouslayer. The dry matter in the mucilage consists of 49 per centpolysaccharides and 22 per cent proteins. The polysaccharidefraction, analysed by ion exchange chromato-graphy, consistsmainly of glucose, arabinose, galactose and uronic acids. Theprotein fraction was analysed by SDS gel dectrophoresis andby high performance liquid-chromatographic separation of theamino acids. The analysis shows a protein pattern very similarto that of the peripheral ligule tissue. It is suggested, therefore,that the material of the external mucilage is secreted by theperipheral ligule cells. The secretional mechanism appears tobe a direct release of polysaccharides and proteins by the tubularcomponents of the ER. There is no indication of secretion viathe Golgi apparatus. Because of its high activity in proteinsynthesis and secretion, it is suggested that the ligule isa vestigial structure, which, in extinct genera, might havefunctioned as a digestive organ. Isoetes lacustris, endoplasmic reticulum, ligule, ultrastructure, Polysaccharide secretion, protein secretion     

Ultrastructure of the Resin Ducts of Mangifera indica L. (Anacardiaceae). 3. Secretion of the Protein-polysaccharide Mucilage in the Fruit

Mango fruit ducts secrete a protein-carbohydrate mucilage inaddition to lipophilic material. This mucilage is secreted inspecial duct regions. Loops of ER elements seem to delimit cytoplasmicportions rich in ribosomes forming pseudo-vacuoles which eventuallybecome bound by a single membrane of ER origin. It is suggestedthat the protein is produced and accumulates in the pseudo-vacuoleswhich become storage bodies. Carbohydrates are added to thecontent of these bodies by Golgi vesicles. The cytoplasm becomesosmiophilic and contracts before disintegration, and the mucilagepasses into the space between plasmalemma and cell wall. Afterthe cell breaks down the mucilage is released into the ductcavity. Mangifera indica L., mango, Anacardiaceae, resin ducts, secretion, mucilage, ultrastructure     

Studies on the Development of Glandular Hairs in Nymphoides peltatum and the Ultrastructure During Their Secretory P

Each glandular hair of Nyrnphoides peltaturn (Gmel.) O. Kuntz consisted of only one row of cylindar cells with secretory function. The hairs originated from the protoderm cells on the adaxial surface of the second leaf primordium from the shoot apex. Cells of the glandular hairs prossessed dense cytoplast during the secretory period, but the vacuoles were very small. There were not only abundant mitochondria, Golgi bodies and endoplasmic reticulum in the glandular hair cells, but also many plasmodesmata. The authors' research indicated that the mucilage was carried to the edge of the cells by the membranous multilamellar bodies and the vesicles from both Golgi bodies and endoplasmic reticulum. The mucilage was secreted extracellularly by either exocytosis or ecrine secretion. The side walls of the glandular hairs swelled because of mucilage mass accumulation in the walls. The mucilage, being tested to be composed of polysaccharides and a trace of protein, played an important role in protecting the development of the vegetative buds of N. peltatum.     

菱的腺毛发育及分泌活动的超微结构研究

菱的腺毛由单列圆筒状细胞组成,具有短暂的分泌功能。它们起源于叶片远轴面、叶柄的表皮细胞以及苗端茎轴、花柄的表皮细胞。处于分泌期的腺毛细胞其胞质浓厚,液泡化程度小,细胞具丰富的线粒体、高尔基体。腺毛丧失了分泌功能后即发育成为表皮毛。粘液物质由高尔基体分泌小泡携带至腺毛细胞侧壁,经胞吐与渗透结合的方式分泌至细胞外,粘液的化学成分主要为多糖。     

The Ultrastructure of Glandular Trichomes ofPhillyrea latifoliaL. (Oleaceae) Leaves

The anatomy and ultrastructure of glandular trichomes at differentdevelopmental stages were investigated inPhillyrea latifoliaL.leaves by transmission electron microscopy and histochemicaltechniques. The trichome consisted of a multicellular secretoryhead, a unicellular stalk and a collecting cell surrounded byepidermal cells and spongy mesophyll cells. There were numerousplasmodesmata across the cell walls of trichome cells, and especiallybetween the stalk cell and the collecting cell. The collectingcell and stalk cell contained few chloroplasts. Mitochondria,elements of the endoplasmic reticulum and small vacuoles wereabundant in the secretory cells. Crystals were present in thesecretory cells and the collecting cell, especially at the matureand senescent stages of trichome development. As the cuticle,which covered the secretory cells, did not show pores or perforations,it is proposed that secretion occurred by accumulation of productsin subcuticular spaces followed by diffusion through the cuticle.Callose accumulation was observed between the stalk cell andthe collecting cell of senescent trichomes, especially in salt-treatedplants. Trichome ontogeny was accelerated in salt-treated plants.Copyright1998 Annals of Botany Company Cuticle;Phillyrea latifolia; secretion; transmission electron microscopy; trichome development.     

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     

Accessory Pollen Adhesive from Glandular Trichomes on the Anthers of Leonurus sibiricus L. (Lamiaceae)

Abstract: Glandular trichomes (ca. 16 per anther) on the anthers of Leonurus sibiricus produce a secretion that, when touched, is liberated at once and becomes sticky when in contact with the air. With successive visitations of the pollinators (species of Bombus in naturalized populations) the number of secretion‐containing glands on each anther diminishes by mechanical rupture. On the pollinators, the secretion mixed with pollen was found adhered to the integument on the parts making contact with the anthers and stigma, mostly on the scape of the antennae. These trichomes are anatomically identical to the glandular scales common in the entire family and are formed by a multicellular cuticle‐bounded structure, with a foot and head. The secretion is accumulated as a milky emulsion under the cuticle, outside the primary cell wall, and is liberated by rupture of the cuticle. The composition possibly differs from what generally distinguishes these glandular trichomes, i.e. volatile oils that give these plants their particular smell. Such volatile compounds are generally assumed to have defensive or attractive functions, different from those observed in this study, which would be strictly mechanical.     

Die Morphologie der Schleimsekretion im Fruchtknoten vonAptenia cordifolia

Zusammenfassung Der Fruchtknoten vonAptenia cordifolia enthÄlt wÄhrend der Samenentwicklung einen proteinreichen Polysaccharidschleim. Verschieden alte schleimproduzierende Placentarpapillen werden einer elektronenmikroskopischen Analyse unterzogen. Kurz vor dem Einsetzen der Schleimproduktion ist das rauhe ER noch spÄrlich entwickelt. Der Golgi-Apparat ist unauffÄllig und wenig aktiv. Zu Beginn der Schleimbildung sind als hauptsÄchliche Strukturkomponenten hypersekretorische Dictyosomen und ER-umschlossene Vakuolen (storage vesicles) zu beobachten. Es wird angenommen, da\ diese Komplexe aus rauhem ER und vermutlich mitèinander verschmolzenen Golgi-Vesikeln die charakteristischen Synthese-Einheiten für den Polysaccharid-Protein-Schleim darstellen, da sie nachweislich neben Polysacchariden auch Proteine enthalten. Membranfusionen zwischen Vesikeln und dem Plasmalemma deuten auf Exocytose-Prozesse unter Beteiligung des Golgi-Apparates hin. Daneben wird eine holocrine Ausscheidung des in den storage vesicles zunÄchst gespeicherten Polysaccharid-Protein-Schleimes bei Degeneration des Protoplasten vermutet.

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Morphology of slime secretion in the seed vessels ofAptenia cordifolia

Summary During seed development the gynaeceum ofAptenia cordifolia produces a mucilage rich in carbohydrates and protein. The mucilage-producing placentary papillae are analyzed in different developmental stages by electron microscopy. Just before mucilage production is started, the rough ER occurs but sparsely. At that time the dictyosomes are inconspicuous and of low activity. When mucilage production commences, one can observe hypersecretory dictyosomes and ER-ensheathed vacuoles (storage vesicles) as the main structural components. It is suggested that the complexes of rough ER and probably fused Golgi vesicles are the synthetizing units of the carbohydrate protein mucilage, since in these complexes both components can be identified cytochemically. Fusion sites of plasmalemma and vesicles indicate processes of exocytosis-probably involving the Golgi apparatus. In addition, a holocrine excretion of the mucilage initially enclosed in the storage vesicles via degeneration of the protoplast is assumed.

     

Ultrastructure and Secretion of Extrafloral Nectaries of Plumeria rubra L.

Extrafloral nectaries situated on the adaxial side of the petiolebase are differentiated into a long head, comprising subepithelialground tissue surrounded by a layer of elongated palisade-likeepithelial cells and a short stalk from the nectary meristem.Many ultrastructural changes occur in epithelial and subepithelialcells of the nectary, from the young to secretory stages, suchas an increase in the amount of cytoplasm rich in mitochondriawith well developed cristae, rough endoplasmic reticulum (rER),smooth endoplasmic reticulum (sER) tubules and Golgi bodies.Plasmalemma invaginations with secretory vesicles occur longthe radial walls. Substantial amounts of secretory materialaccumulate in the gap between the radial walls and subcuticularspace, probably carried by the secretory vesicles from the cytoplasmat the secretory stage. Before cessation of secretion the cytoplasmbecomes vesiculated and the volume of the vacuome increases.At the post secretory stage, cytolytic processes and death ofcells occur. The subepithelial cells attain their maturity priorto epithelial cells. Histochemical localization reveals thepresence of lipids, proteins and insoluble polysaccharides withinthe epithelial cells and in the secretory material depositedin the subcuticular space as well as the gap between the radialwalls of the epithelial cells and outside the cuticle. Fine structure, nectary, Plumeria rubra, granulocrine secretion     

DEVELOPMENT,STRUCTURE AND FUNCTION OF SECRETORY TRICHOMES IN PSYCHOTRIA BACTERIOPHILA (RUBIACEAE)

Mucilage-secreting dendroid trichomes develop from the adaxial epidermis of young stipules surrounding the shoot apex. Each trichome consists of a multicellular stalk from which radiate many branch cells. The trichome has no cuticle and the branch cell walls distally are loose cellulosic frameworks. Dictyosomes produce vesicles whose products are secreted through the plasma-lemma and cell wall. Enlarged portions of the ER are frequently associated with dictyosomes and may be part of the system for synthesis and transport of secretion products. Bacteria, which later occur in leaf nodules, are present in the mucilage surrounding trichomes and young leaves. The latter develop stomata through which the bacteria enter. As stipules and leaves grow out of the apical region, the secretory trichomes degenerate and are replaced by non-secretory ones.     

Cytochemical and Morphological Evidence for the Involvement of the Plasma Membrane and Plastids in Mucilage Secretion in Aloe arborescens

Most of the volume of Aloe arborescens leaves (volumetric density= 0.68±0.08) is occupied by a mucilage tissue. The mucilagesof Aloe species are glucomannans of great medical and pharmaceuticalimportance. An electron microscopical and histochemical studywas carried out, following the development of the tissue andsecretion processes. In contrast to other reports on polysaccharide secretion inplants, no observable activity was found in the Golgi apparatus.In the young stages of leaf development, positive histochemicalstaining for polysaccharides and structural changes were foundmainly in the plastids. In the mature leaves the mucilage cellsremain alive and intact and secretory activity appears to becentered in the plasma membrane. Aloe arborescens, cytochemistry, mucilage, plasma membrane, plastids, secretion     

Pollen--stigma Interaction in the Leguminosae: the Secretory System of the Style in Trifolium pratense L.

The canal that traverses the upper part of the style of Trifoliumpratense is derived lysigenously. The core tissue of the veryyoung style consists of elongated cells similar to those ofthe transmitting tissue of solid-style families such as theSolanaceae; as the style matures, these cells separate to formthe canal, which receives secretions both from the core tissueand the inner wall cells. The early secretion of proteins intothe intercellular spaces is associated with the presence ofparamural bodies (lomasomes) in the adjacent cells. In the cellsin the immediate vicinity of the canal, vesicles, probably derivedfrom the Golgi system enlarge during later development and accumulatea protein-carbohydrate content, which is later passed into thecytoplasm where it forms densely packed fibrillar nodules. Withthe dissolution of the cell membranes, this material is passedinto the canal, where it is progressively diluted by continuedingress of water until the cavity reaches its final volume. Leguminosae, Trifolium pratense L., pollen—stigma interaction, self-incompatibility, stylar secretion, protein secretion     

Anatomical studies of the secretory structures: Glandular trichomes and ducts, in Grindelia pulchella Dunal (Astereae, Asteraceae)

The ultrastructure of the glandular trichomes and secretory ducts of Grindelia pulchella was studied. Plastids, mitochondria and endoplasmic reticulum are involved in the secretory process of both, trichomes and ducts. A special tissue with “transfer cells” is associated with the duct epithelial cells. The secretion is produced in the transfer cells and then is transferred to the duct epithelial cells where it accumulates in the vacuoles. The occurrence of cavities within the cell walls of the trichome cells and duct epithelial cells is described. The secretion is accumulated between the cell wall and the cuticle of these cells. When the cuticle is broken the secretion is released. We conclude that granulocrine secretion operates in this species.     

The Structure and Cytochemistry of the Pistil of Sternbergia lutea (Amaryllidaceae)

Studies were carried out on structural and cytochemical aspectsof the pistil of Sternbergia lutea (L.) KerGawl. The stigmais of the wet papillate type; the papillae are unicellular andare arranged densely around the rim of a funnel-shaped stigma.The stigma exudate is limited and is confined to the bases ofthe papillae and the inner lining of the stigma. The papillaeare smooth in the distal part and are covered with intact cuticle-pelliclelining. The cuticle is disrupted at places towards the baseof the papillae releasing the exudate. The exudate is rich inpectins and other polysaccharides but poor in proteins and lipids.The papillae show dense cytoplasmic profiles with extensiveendoplasmic reticulum (ER), abundant mitochondria, polyribosomesand active dictyosomes. The style is hollow. The stylar cavityis surrounded by two to four layers of glandular cells. In theyoung pistil the canal is lined with a continuous cuticle, butin the mature pistil the cuticle becomes disrupted and the canalis filled with the secretion produced by the cells of the surroundingglandular tissue. Ultrastructurally, the cells of the glandulartissue are very similar to the stigmatic papillae. The innertangential wall of the cells bordering the canal is uniformlythicker than other walls. The secretion in the stylar canal,as well as the intercellular spaces of the glandular tissue,stain intensely for pectins and polysaccharides but poorly forproteins and lipids. Pollen tubes grow through the stylar canal.Structural and cytochemical details of the pistil of Sternbergiaare compared with other hollow-styled systems. Pistil, Sternbergia lutea (L.) Ker-Gawl., stigma and style, structure and cytochemistry