Leaf Nodule Development in Psychotria kirkii Hiern. (Rubiaceae)

The initiation, development and structure of the leaf nodulesof the Rubiaceous shrub Psychotria kirkii Hiern. has been studiedin detail at the ultrastructural level. Bacteria, maintainedin the shoot tip in the secretions from dendroid colleters,invade the substomatal chamber of stomatal pores which formprecociously on the abaxial leaf surface. Proliferation of theepidermis around the pore pushes the bacterial cavity deep intothe lamina, thus forming a small internal nodule. Endophyte-mediatedschizogeny of the cells surroundng the nodule causes it to expandwhile at the same time giving rise to an interconnected reticulumof invasive host cells which are involved in metabolite exchangebetween microoganisms and host plant. Bacterial morphology changesafter entry of the microsymbiont into the host plant and, bythe time the nodule is mature, the bacteria exhibit distinctpleomorphism. Senescent nodules are shown to accumulate lipidand starch. The developmental process is discussed in the lightof existing information on this symbiosis. Psychotria kirkii, leaf nodule development, symbiosis, ultrastructure     

The occurrence of calyx nodules inPsychotria spp. (Rubiaceae)

Summary The occurrence and structure of calyx nodules in the flowers of two leaf nodulated rubiaceous speciesPsychotria punctata Vatke andPsychotria kirkii Hiern. has been described for the first time at the ultrastructural level. Bacteria, resident in colleter-secreted mucilage in the space between calyx and corolla, invade stomatal pores which develop on the calyx protoderm. The bacteria proliferate in the substomatal cavity and then invade the calyx mesophyll. This invasion is most pronounced inP. punctata where the bacteria even penetrate and enter the cells of the vascular tissue. Although no sheath forms around the calyx nodules, the calyx mesophyll cells surrounded by the bacteria become identical in shape, size and secretory function to the invasive mesophyll cells of leaf nodules. The functional and evolutionary significance of calyx nodulation is discussed.     

Colleter morphology in Pavetta, Neorosea and Tricalysia (Rubiaceae) and its relationship to the bacterial leaf nodule symbiosis

Buds removed from herbarium specimens were processed for light microscopic examination of colleter form and anatomy. Most Pavetta species have dendroid colleters and bacterial leaf nodules, a correlation also found in an earlier study of Psychotria. Colleters of Neorosea , another genus with leaf-nodulated species, are more like the standard rubiaceous type that predominates in the family except that they have irregular, bulging epidermal cells. Tricalysia , a nodule-free genus closely related to Neorosea , shows a range of colleter form from standard to dendroid, and some species have the Neorosea type of colleter. Such morphological correlation between the bacterial leaf nodule symbiosis in Pavetta and Psychotria and dendroid colleters may indicate a chemical change in colleter secretion. Apart from their involvement with the bacterial symbiosis, rubiaceous colleters have now been shown to vary sufficiently in certain taxa to be considered as additional useful taxonomic characters.     

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.     

Colleter types in Rubiaceae, especially in relation to the bacterial leaf nodule symbiosis

Vegetative buds were removed from herbarium specimens, and a small number of living plants, of 116 species of 111 genera of Rubiaceae and processed for light-microscopic examination of colleter form and anatomy. The results were combined with those of earlier studies. Most colleters in this family are of the 'standard type' described by others. Colleters are, however, lacking in eight genera. dendroid colleters occur in six genera. Two other genera have an intermediate type of colleter. Three genera have bacterial leaf nodules; dendroid colleters are not restricted to these genera, but their pattern of occurrence suggests a causal relationship between bacterial infection and evolution of dendroid colleters. Coupled with the geographic distribution of nodulared species within these genera, this pattern indicates that Pavetta became infected very early in its evolutionary history, Psychotria relatively later, and Neorosea most recently.     

Structural, Functional and Phylogenetic Aspects of the Colleter

This article surveys the structural, functional and phylogeneticsignificance of colleters in different dicotyledonous families.Colleters are multicellular secretory structures attached tothe stipule, petiole, lamina, bract, bracteole, calyx and corolla.Colleters are grouped into standard (S), dendroid (D) and brush-like(B) types on the basis of their morphology and structure. Dand B-type colleters occur in certain members of Rubiaceae thatalso have bacterial leaf nodules. Besides the normal structure,epithelial hairs, thin-walled subepidermal cells, laticifersand vasculature are present in many colleters of Apocynaceae.It is probable that the colleter functions to protect the developingmeristem by secreting a viscous fluid. Exudate of D-type colletersare mucilaginous, providing the substrate necessary for thenutrition of endophytic bacteria. Colleter, structure, phylogeny     

Trichome forms in Ardisia (Myrsinaceae) in relation to the bacterial leaf nodule symbiosis

Symbiotic leaf-nodule bacteria in nodulated members of Rubiaceae live in mucilage secreted by colleters located on stipules within buds. These differ from colleters on most nodule-free species. This study was undertaken to examine buds of Ardisia and the related monotypic Amblyanthus of Myrsinaceae to see if nodulated species had secretory structures dissimilar from those of nodule-free species. Buds removed from herbarium specimens (61 species) and live plants (3 species) were paraffin-sectioned. Diverse trichome forms occur, including dimorphism between adaxial and abaxial trichomes in some species. Species within each subgenus were arranged according to trichome form: peltate scale, irregularly capitate, capitate, sessile capitate, bicellular capitate, and uniseriate. Only seven Ardisia species (all in subgenus Crispardisia, widely assumed to have bacteria in marginal leaf nodules of all 30 species) have short-lived trichomes bearing one or more elongate, swollen, distal cells that appear to be secretory cells. These trichomes are analogous to the dendroid or brushlike colleters of nodulated Rubiaceae. Druses occur in most subgenera and this appears to be by far the most predominant crystal type m Ardisia.     

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.     

Development, Structure and Cytochemistry of Resin-secreting Colleters of Gardenia gummifera (Rubiaceae)

The structure, ontogeny and cytochemistry of stipular colletersof Gardenia gummifera are described. The repeated division ofa few adaxial protodermal and subprotodermal cells of the youngstipule and the subsequent elongation by the latter resultsin the formation of a standard-type colleter. A developed colleterhas outer palisade-like glandular cells covered by a continuouscuticle and inner layers of elongated columnar cells. Enzymelevels in the palisade-like epidermal cells in the colletersindicate their active role in secretion. The secretion is resinous,occurs in the young colleters, and accumulates between the cellsand the cuticle. The cuticle later ruptures and releases thesecretory product, which drenches the young shoot apex. Gardenia gummifera L, stipule, colleters, resin, ontogeny, cytochemistry     

Histochemistry and Senescence of Colleters of Allamanda cathartica (Apocynaceae)

Allamanda have colleters on the adaxial bases of petiole, bractsand bracteoles. Each colleter is finger shaped, differentiatedinto a long head on a short stalk. Vasculature to the colleteris absent but branched laticifers are present among the centralcells. Secretion of the colleter contains glucose and rhamnose.The major elements present in the exudate are Na, Fe and Zn.Starch, protein and lipids were identified, with lipids themajor component. Petiolar colleters are persistent and becomestiff due to the lignification occurring in the walls of epithelialand central cells. Because of over-lignification in the centralcells, the cell lumen is highly reduced. Allamanda, colleter, laticifer, secretion, senescence     

The Extrafloral Nectaries of Cotton: I. FINE STRUCTURE OF THE SECRETORY PAPILLAE

The fine structure of the secretory papillae of the extrafloralnectaries of cotton (Gossypium hirsutum) is described. The cellscontain a dense cytoplasm with the rough endoplasmic reticulumbeing particularly prominent. The cuticle covering the papillaehas a typical two-layered appearance and is detached from thewall in secretory cells. With maturity, the lateral walls ofthe stalk cell at the base of each papilla become impregnatedwith cuticle-like electron-opaque material. The frequency anddistribution of plasmodesmata have been estimated in all wallsof the papillae. The periclinal walls are traversed by numerousplasmodesmata (about 16 per µ m² in the distal wall ofthe stalk cell) which, in general, change from a simple to amore complicated structure during nectary development. The resultsare discussed in relation to the role of the ER in nectar secretionand are considered to support the view that pre-nectar followsa symplastic pathway from the phloem to the secretory cells. Key words: Gossypium hirsutum, Secretory papillae, Nectary     

Ontogenesis secretion and senescence of Tocoyena bullata (Vell.) Mart. (Rubiacaeae) colleters

Colleters are secretory structure present on many families including Rubiaceae. Particular characteristics have been described about colleters secretory cells, however senescence process are still under debate. Tocoyena bullata (Vell.) Mart. (Rubiaceae) shoot apex were collected at Jardim Botânico do Rio de Janeiro, RJ/Brazil. Stipules were separated and fragments were fixed in 2.5% glutaraldehyde and 4.0% formaldehyde in 0.05 m sodium cacodylate buffer, pH 7.2, post fixed in 1.0% osmium tetroxide in the same buffer, dehydrated in acetone, critical‐point‐drying, sputtered coated and observed. For light microscopy fragments were fixed and dehydrated, infiltrated with historesin and stained with 1% toluidine blue. For transmission electron microscopy, the samples were infiltrated with Epoxi resin. Colleters are present on stipule adaxial surface. On the beginning of development, these structures are recognized as small projections. Later on, colleters differentiated and secrete by cuticle rupture. The colleters senescence occurs in a concomitant and indissoluble way of programmed cell death. Ultrastructural analyses during the process strongly suggest the senescence is based on a non‐autolitic programmed cell death. T. bullata colleters, present at stipule abaxial surface are cylindrical secretory structures. Colleters secretory cells originated as stipule projections; differentiate; secrete and senesce by programmed cell death. The secretion and the cell dead occurs in a concomitantly and indissoluble way.     

Glandular Trichomes of Fagonia L. (Zygophyllaceae) Species: Structure, Development and Secreted Materials

The glandular trichomes ofFagoniaconsist of one secretory celland a multicellular stalk, which develops by division, elongationand elevation of epidermal cells. The latter become seperatedfrom the mesophyll and a subepidermal chamber is formed. Thelength of the stalk, which differs in the various species orvarieties is determined by the number of cell divisions and/or the extent of cell elongation. Although the basic morphologyand development of the trichomes of the species and varietiesexamined are similar, two types of mature trichomes can be distinguished:one occurs in the two examined varieties ofF. mollisand thesecond inF. glutinosaandF. arabica. The secretory cells of thesecond type possess a very thick wall and bear a porous cupuleon their top. Histochemical tests revealed that the sticky substancesecreted by the secretory cells contains mainly polysaccharidesand lipophilic compounds. The secreted material exhibits autofluorescence.InF. mollisvar.hispidachanges in the amount and shape of thefluorescent material inside the secretory cell, during the courseof development, have been observed. The contribution of theglandular trichomes inFagoniaspecies to survival in hot desertconditions is discussed. Fagonia; glandular trichomes; subepidermal chamber; secreted material; adaptation to desert conditions; stalk; fluorescence     

Pollen-Stigma Interaction in the Leguminosae: the Organization of the Stigma in Trifolium pratense L.

The intact stigma of Trifolium pratense possesses a smooth receptivesurface fringed by a few ranks of brush hairs. This surfaceis ensheathed by a thin (75–100 nm) but highly impermeablecuticle, which encloses four to five ranks of secretory cellsimmersed in their secretory products. Experimental single-grainpollinations show that pollen cannot become hydrated or germinateon the intact surface. The cuticle is ruptured when the floweris tripped; the secretion is released, and captured pollen-selfor cross-can then germinate. As in other papilionoid Leguminosae,this mechanism provides a guard against premature selling. Thesecretory cells are elongated; they remain in communicationthrough persistent pit-fields as the intercellular spaces fillwith secretion product. The secretion forms a lipid-rich emulsion,with a mucilaginous aqueous phase which reacts cytochemicallyfor protein and carbohydrate and has esterase activity. Duringthe early development of the stigma head, the cells possessa fine-structure appropriate to their secretory function, withabundant ribosomal and smooth endoplasmic reticulum, stratifiedor in the form of ramifying and anastomosing tubules, numerousmitochondria and a well developed Golgi system. Lipid globuli,partly invested in endoplasmic reticulum, are abundant in theyoung cells, but there is as yet no indication of how the lipidis transferred to the intercellular spaces during the secretoryperiod. As the stigma matures, the secretory cells become moribund. Leguminosae, Trifolium pratenseL., pollen-stigma interaction, self-incompatibility, autofertility, stigma secretory system, lipid secretion, cuticle permeability     

The Floral Nectaries of Hibiscus rosa-sinensis: 1. Development of the Secretory Hairs

Floral nectaries of Hibiscus rosa-sinensis occur on the lowerinner side of the fused sepals and each one consists of numerous(50000–55000) secretory hairs, occupying a cylinder-likezone completely lining the inner side of the sepals. Each hairoriginates from a single protodermal mother cell and, at maturity,it is built up of a basal cell, a stalk, 35–40 intermediatecells and a tip secretory cell. Development of protodermal cellsinto secretory hairs is asynchronous, the first cells to initiatedevelopment being those situated in the lowermost part of thecylindrical zone, and development progressing upwards. Volume increase of protodermal mother cells initiating developmentis accompanied by cell polarization manifested by organelledisplacement towards the apical region. Secretory hairs areformed through a sequence of transverse and, later on, anticlinaldivisions. Divisions of apical cells are preceded by well definedpre-prophase microtubule bands, which foreshadow the plane ofthe forthcoming division and predict with accuracy the sitesof parental walls where the new cell plate fuses at cytokinesis. Stalks consist of either one or two cells. Two-celled stalksoccur in 40 per cent of secretory hairs and derive from a transversedivision of one stalk cell; the wall formed is always depositedparallel to the proximal and distal walls, but never to thelateral ones. The significance of this mode of division is discussedin relation to the fact that lateral walls are entirely impregnatedwith a cutin-like material that blocks apoplastic movement ofsolutes. Hibiscus rosa-sinensis, nectaries, development, preprophase microtuble bands, stalk cells     

Leaf colleters in Tontelea micrantha (Celastraceae,Salacioideae): Ecological,morphological and structural aspects

The colleter secretion can be useful to protect plants of Cerrado (Brazilian savanna) biome during the long and pronounced dry season. This study describes the presence of colleters in Tontelea micrantha and represents the first record of these structures in Celastraceae. To investigate colleter structure and their secretory processes, young leaves were collected, fixed, and processed according to conventional techniques for light, and electron microscopy. Colleters were observed at the marginal teeth on the leaf. They produce mucilaginous secretions that spread over the leaf surface. After secretory phase, colleters abscise. The secretory epithelium is uniseriate and composed of elongated cells whose dense cytoplasm is rich in organelles. The ultrastructure of the secretory cells is compatible with the pectin-rich secretion. Observations of the young leaves surface revealed the presence of superficial hydrophilic secretion films that appeared to have the function of maintaining the water status of those organs.     

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     

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     

Anatomical structure and secretion compounds of colleters in nine Ilex species (Aquifoliaceae) from southern South America

In order to clarify whether the structures observed at the base of the petiole of the genus Ilex are colleters resulting from stipules, the anatomy, vascularization and secretions of these supposed glandular structures were analysed in nine species. This is the first report of colleters in Ilex. Stipular colleters replace the stipules in all species studied and are characterized by the presence of vascular traces. In addition to the stipular colleters, three other types of colleter were distinguished: standard and lachrymiform colleters found on the leaf teeth or crenations, and sessile colleters found on the margins of the floral bracts. Their basic structure consists of a central core of parenchymatous cells surrounded by one layer of palisade secretory epidermal cells. Histochemical tests were also performed on secretions; proteins were found in the secretions studied, but glucose was not. The glandular origin of the stipular colleters is confirmed on the basis of their position, secretions and anatomy. Analyses of the colleter‐secreted proteins distinguished two different groups of Ilex species. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160 , 197–210.