Development and Structure of Primary Secretory Ducts in the Stem of Commiphora wightii (Burseraceae)

Development of gum-resin ducts, sites of resin synthesis inthe epithelial cells and elimination of resin from the protoplasthave been studied in the stem of Commiphora wightii. Formationof an intercellular space amongst a group of densely stainedprocambial cells signals the initiation of a duct. It widensby anticlinal divisions of the epithelial cells and also bytheir further separation along the radial walls. The numerousplastids with varying morphological shapes have an electrondense matrix. Starch granules present in the amyloplasts showevidences of exocorrosion. Mitochondria in the epithelial cellsof developing and mature ducts have well developed, swollencristae. Osmiophilic material originates in association withthe golgi-derived vesicles at its maturing face. It is alsoobserved in close association with plastids, mitochondria andvacuoles, thereby plausibly involving them in the process ofresin formation. The resinous material is eliminated from thecytoplasm by vesicles enveloped by plasmalemma prior to theirdischarge into the apoplast. Myelin-like multilamellate structuresobserved along the inner tangential wall may aid in the secretionof resin across the wall. Commiphora wightii, primary secretory ducts, epithelial cells, ultrastructure, gum-resin secretion     

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 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.     

Effects of Wounding and Fungus Inoculation on Terpene Producing Systems of Maritime Pine

Cheniclet, C. 1987. Effects of wounding and fungus inoculationon terpene producing systems of maritime pine.—J. exp.Bot. 38: 1557–1572. The qualitative and quantitative changes occurring in terpenecompounds (resin acids and mono- and sesquiterpene hydrocarbons)in phloem and xylem tissues after wounding and after inoculationwith a beetle-associated fungus have been compared on maritimepines from the same clone. Simultaneously, structural eventsoccurring in phloem have been observed, especially those concerningresin ducts and cell systems associated with terpene secretion. The terpene accumulation resulting from wounding is weak andremains limited to a short period and to a restricted tissuearea. In contrast, mycelium inoculation results in an exponentialenrichment (x 60 after 28 d in proximal phloem) as the enrichedzone enlarges, indicating a diffusive process. In wounded phloem the neosynthesis of terpenes takes place inreactivated or supernumerary resin ducts which display structuralfeatures similar to those of active secretory cells in normalconditions. In fungus-moculated phloem, secretory ducts are first reactivated,but soon undergo a lysis, and are no longer functional whenthe rate of terpene accumulation is increasing quickly. Theformation of terpene compounds is delocalized and occurs innon-specific intra- or extracellular spaces formed by the lysisof cell contents and/or cell walls. The synthesis probably resultsfrom the random meeting of enzymes, which can be synthesizedin living parenchyma cells or released by lysis of structuresinvolved in physiological terpene secretion (leucoplasts andendoplasmic reticulum), and of the corresponding substrates,which may be provided by degradation of cell contents (polyphenolics,hpids) or by the sieve flow. Key words: Terpene secretion, wounding, fungus inoculation     

Structure of the Needles in the Early Phases of Development in Pinus ponderosa P. et C. Lawson with Special Reference to Plastids

Before they emerged from the fascicular sheath, the tissuesof young needles of Pinus ponderosa P. et C. Lawson alreadyshowed some characteristics typical of mature needles. The organelles,particularly the plastids, had undergone different development.The plastids in different types of cell varied in their ultrastructureand in association with the endoplasmic reticulum (ER). A sheathof ER was observed around the amoeboid plastids in epidermalcells, epithelial cells of resin ducts and maturing transfusiontracheids whereas there was no ER sheath around the young mesophyllchloroplasts, the fusiform chloroplasts in some transfusiontracheids and the proplastids in xylem and phloem cells. Thecontent of chlorophyll (a+b) was 0·85 g kg^-1 dry matterand chlorophyll a/b ratio was 2·70. The needles may becomephotosynthetically active whilst still within the fascicularsheaths.Copyright 1993, 1999 Academic Press Pinus ponderosa, ponderosa pine, needle structure, needle ontogeny, plastids, endoplasmic reticulum     

The Digestive Glands of Pinguicula: Structure and Cytochemistry

The digestive glands of the carnivorous genus Pinguicula havethree functional compartments, (a) a basal reservoir cell, (b)an intervening cell of endodermal character and (c) a groupof secretory head cells. The gland complex is derived from asingle epidermal initial. The reservoir cell, which is richin Cl^– ions, is highly turgid before discharge; it islinked by plasmodesmata to the surrounding epidermal cells,and is ensheathed by a pectin-rich inner wall layer. The endodermalcell is bounded by a Casparian strip to which the plasmalemmais tightly attached; it contains abundant storage lipid andnumerous mitochondria. The head cells of the developing glandhave labyrinthine radial walls of the transfer-cell type, theingrowths being composed of pectic polysaccharides. The boundingcuticle is discontinuous, although lacking well-formed pores.Mitochondria are numerous, with well-developed cristae; theplastids are large and ramifying, and invested by ribosomalendoplasmic reticulum. Dictyosomes are sparse, and where theyoccur, are associated with coated vesicles. Ribosomal endoplasmicreticulum is moderately abundant in the head cells, and so alsoare free ribosomes. Optical and electron microscopic localizationmethods indicate that the digestive enzymes are synthesizedin the head cells and transferred both into the vacuoles andinto the walls. There is no evidence of a granulocrine modeof secretion, and the transfer seems to be initially by directperfusion through the plasmalemma. During the final phase ofmaturation of the head cells they suffer a form of autolysis,vacuoles, cytoplasm and wall becoming confluent as all of themembranes of the cell undergo dissolution. The gland head isthus, in effect, simply a sac of enzymes at the time of theultimate discharge. Pinguicula, carnivorous plant, insectivorous plant, enzyme secretion, digestive gland     

Anatomy of Secondary Phloem of Glyptostrobus in Relation to Its Systematic Position

A anatomical characters of secondary phloem in Glyptostrobus pensilis (Staunt.)Koch were observed by means of both light and scanning electron microscopy(SEM). The secondary phloem is composed of axial and radial systems. In the axial systems, the phloem consists of sieve cells, phloem parenchyma cells, albuminous cell and phloem fibers. In the radial systems, it consists of phloem rays. The alternate arrangement of different cells in cross section results in tangential bands. The sequence of radial arrangement follows the pattern of sieve cells, phloem parenchyma cells, sieve cells and phloem fibers, sieve cells. Many crystals of calbium oxalate are embedded in the radial walls of seive cells. The phloem fibers are of only one type. The phloem rays are homogeneous, uniseriate. According to the anatomical characters of secondary phloem of Glyptostrobus pensilis (Staunt.)Koch and comparison with the other genera of Taxodiaceae, Glyptostrobus, Metasequoia and Taxodium have close relationships.     

Ultrastructure of Gum-resin Ducts in Cashew (Anacardium occidentale)

In Anacardium occidentale L., the gum-resin ducts in the primaryphloem of the stem develop schizogenously. Appearance of anintercellular space amongst the densely stained cells signalsthe initiation of a duct. During this process, the middle lamella,appears in places, as an electron-opaque area. The epitheialcells bordering the duct are oval and have convex inner tangentialwalls which are without any plasmodesmata, although they areabundant on radial and outer tangential walls. The cytoplasmof the epitheial cells is rich in rough endoplasmic reticulum(ER), free ribosomes, polysomes, mitochondria with swollen cristae,plastids with occasional osmiophilic inclusions, dictyosomesand vesicles. Osmiophilic material has been observed in thedilation of the cisternae of dictyosomes and also in vesicles.Sometimes, the osmiophilic material aggregates and forms largemembrane-bound globules. The globules fuse with the plasmalemmaat the inner tangential wall, and presumably the contents aredeposited in the space between the protoplast and the wall.This material passes through the loose matrix of the wall intothe duct. Some of the epitheial cells of the mature duct show‘dark’ cytoplasm, degraded organelles and occasionallyhigh vacuolation; ultimately they are lysed and their remainscollect in the duct. Anacardium occidentale L., cashew, gum-resin ducts, epithlial cell, dictyosome, osmiophilic material, ultrastructure     

Ontogeny of anastomosed laticifers in the stem apex of <Emphasis Type="Italic">Hancornia speciosa</Emphasis> (Apocynaceae): a topographic approach

Latex is a complex plant secretion with both ecological and economic importance. There is little information currently available on the cytological aspects of the ontogenesis of anastomosed laticifers, the ducts originating through the lysis of adjacent cell walls. Hancornia speciosa is a tree typical of the Cerrado (neotropical savanna) biome. Its latex has medicinal value and is also used to produce rubber. The ontogenesis of its laticifers and the process of latex synthesis are described here. Structural, cytochemical, and ultrastructural analyses of the stem apex and phytochemical analyses of the latex were performed. Laticifer ontogenesis begins early in promeristem cells and subsequently extends through the procambial region. The laticifer precursor cells demonstrate intense metabolic activity, evidenced by starch accumulation and the proliferation of mitochondria, dictyosomes, endoplasmic reticulum, and ribosomes—resulting in the thickening of the cell walls and accumulations of oil droplets in the cytoplasm and fibrous materials in the vacuoles. The ontogenetic process culminates with the partial dissolution of adjacent cell walls and the collapse of the cytoplasm, giving rise to anastomosed laticifers distributed throughout the phloem and adjacent regions of the cortex and medulla. The latex itself is composed of terpenes, mucilage, proteins, alkaloids, and organelle residues that form an emulsion. Laticifer development takes place in three phases: (1) the formation of the emulsion in the promeristem, (2) anastomosis and the collapse of the cytoplasm in the distal region of the procambium, and (3) the maturation of laticifers and latex storage in a central vacuole in the proximal region of the procambium.     

水松的次生韧皮部解剖及其系统位置的讨论

在光学显微镜和扫描电子显微镜下观察,水松茎次生韧皮部的主要特征为：韧皮部由轴向系统和径向系统组成。轴向系统由筛胞、韧皮薄壁组织细胞、蛋白细胞和韧皮纤维组成,径向系统由韧皮射线组成。在横切面上,轴向系统的各组成分子以单层切向带交替有规律的排列,其排列顺序为：筛胞－韧皮薄壁组织细胞－韧皮纤维-筛胞。筛胞的径向壁上嵌埋有草酸钙结晶,韧皮纤维仅一种类型,韧皮射线同型、单列。根据水松茎次生韧皮部的解剖研究,并与杉科其它各属的有关资料进行比较,我们认为：水松属与水杉属和落羽杉属有较近的亲缘关系。     

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     

Unusual Network of Internal Phloem in the Pod Mesocarp of Cowpea [Vigna unguiculata (L.) Walp. (Fabaceae)]

A mycelium-like network of internal phloem was observed in theinner mesocarp of the lateral pod walls of the fruit of certaingenotypes of cowpea [Vigna unguiculata (L.) Walp.] In the cultivarVita 3, the network consists of single, or rarely double, strandsof sieve elements and associated phloem parenchyma, orientedmainly parallel with the fibres of the adjacent endocarp, andstretching marginally beyond the sheets of fibres to connectabove and below with the outermost phloem of the longitudinalstrands of the dorsal and ventral sutures of the fruit. Theinternal phloem network does not relate conformationally to,or interconnect with the conventional (xylem+phloem) vasculatureof the mid mesocarp of the pod wall. In Vita 3, sieve elementsdifferentiate in the internal phloem after those in the majorveins of the pod, but before the presumptive endocarp fibrescommence wall thickening. The pod walls of twenty-one otherspecies of legumes proved negative for internal phloem, whileof nine varied genotypes of cowpea examined, six proved positive,three negative for the trait. Presence of internal phloem incowpea is not always associated with presence of endocarp fibresor necessarily with large fruits with large seeds. Possiblefunctions suggested for the phloem network are to provide assimilatesfor fibre wall thickening or to transport solutes to or fromsites of temporary storage in the fleshy inner layers of thepod wall. Internal phloem, legume fruit, translocation, mesocarp, pod wall, Vigna unguiculata, cowpea     

Anatomy of the vegetative organs and secretory structures of Rhaponticum carthamoides (Asteraceae)

Roots, stems, rhizomes and leaves of Rhaponticum carthamoides (Willd.) Iljin (a Siberian adaptogenic plant, originating from the Altai and Saian Mountains) of different ages were investigated by means of light and electron microscopy. Schizogenous secretory reservoirs occurred in every organ, and were located within the secondary xylem (adventitious roots and rhizome of young plants), at the interface of endodermis/cortical parenchyma (roots and hypocotyl), along phloem and primary xylem (older rhizome), around the vascular bundles (inflorescence stem, petiole and leaf midrib veins) and along phloem (cotyledonary and leaf veins). At the interface of endodermis/inner parenchyma, secretion accumulated in the intercellular spaces prior to the formation of proper epithelial cells. The secretion as observed by transmission electron microscopy comprised three components: soluble (i.e. absent from sections; probably phenolic), insoluble and strongly osmiophilic (probably phenolic) and insoluble, moderately osmiophilic (probably lipidic). Numerous osmiophilic oil droplets, similar to the lipidic secretion inside the reservoirs, local proliferation of rough endoplasmic reticulum and numerous multivesicular bodies characterized epithelial cells in all organs. Leucoplasts (in subterranean organs) with osmiophilic inclusions and peroxisomes with crystalloid inclusions were specific for parenchyma cells. Peltate glandular hairs were formed on leaf blades.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 207–233.     

Intracellular Transport and Elimination of Resin from Epithelial Duct-cells of Pinus halepensis

The ultrastructure of the resin secreting cells of root ductsof young Pinus halepensis seedlings was studied. It is suggestedthat the endoplasmic reticulum (ER) in addition to taking partin resin synthesis also plays a role in transporting the resinfrom the plastids, mitochondria and nuclear envelope to theplasmalemma. By fusing with the plasmalemma the ER releasesthe resin to the outside of the protoplast. The resin producedin the ground cytoplasm and by the Golgi apparatus seems tobe eliminated by plasmalemma invaginations. Pinus halepensis, resin secretion, root ducts, endoplasmic reticulum     

The Anatomy of Contractile Roots in Eucomis L'H?rit

The anatomical changes in the different tissues during rootcontraction were studied in two species of Eucomis (Liliaceae).It is evident that the shortening of the root is brought aboutby radial and tangential broadening and longitudinal shorteningof the perivascular cortical cells. This tissue is hence calledcontractile parenchyma. As a result of this contraction thecell walls of the exodermis, endodermis, pericycle, phloem andpith become buckled longitudinally while the annular and spiralthickenings of the xylem are pressed together.     

Development, Histochemistry and Ultrastructure of Gum-resin Ducts in Commiphora mukul Engl.

Gum-resin ducts are present in the primary and secondary phloemof Commiphora mukul Engl. The important gum-resin, known commerciallyas ‘guggul’, is secreted and collected in ductswhich develop schizogenously. The duct initials have dense cytoplasm,large nuclei, increased cytoplasmic RNA and proteins. The lumenof newly-formed ducts widens accompanied by anticlinal divisionsand subsequent tangential elongation of epithelial cells. Histochemicaltests reveal that the epithelial cells have apparently largeamounts of proteins, cytoplasmic RNA, and DNA in the nucleus.Lipid globules are also present in these cells. Epithelial cellwalls in contact with the duct are thin and of a loose fibrillarmesh. Microtubules, randomly oriented in the epithelial cellsare always parallel and adjacent to the wall. The cytoplasmis rich in ribosomes, endoplasmic reticulum, mitochondria, plastidsand vacuoles containing osmiophilic substances. At the peripheralregion of the duct, electron-transparent bodies containing densely-stainedmaterial are present close to the tangential wall.     

Wound-induced traumatic resin duct development in stems of Norway spruce (Pinaceae): anatomy and cytochemical traits

Wounding of Norway spruce by inoculation with sterile agar, or agar containing the pathogenic fungus Ceratocystis polonica, induced traumatic resin duct formation in the stem. Visible anatomical responses occurred in the cambium 6-9 d post-inoculation. Near the inoculation site cellular proliferation, polyphenolic accumulation, and lignification were induced as a wound reaction to seal the damaged area. Five centimetres from the inoculation site cells in the cambial zone swelled and divided to form clusters. By 18 d post-inoculation, these cells began to differentiate into resin duct epithelial cells surrounding incipient schizogenous lumens. Mature axial traumatic ducts appeared by 36 d as a row of ducts in the xylem centripetal to the cambium. The ducts formed an interconnected network continuous with radial resin ducts. Parenchyma cells surrounding the ducts accumulated polyphenols that disappeared as the cells differentiated into tracheids. These polyphenols appeared to contain fewer sugar residues compared to those accumulating in the secondary phloem, as indicated by the periodic acid-Schiff's staining. The epithelial cells did not accumulate polyphenols but contained immunologically detectable phenylalanine ammonia lyase (EC 4.3.1.5), indicating synthesis of phenolics as a possible resin component. These findings may represent a defense mechanism in Norway spruce against the pathogenic fungus Ceratocystis polonica.     

Distribution of leaf assimilates in the stem of Picea abies L.

Summary Autoradiographic and microautoradiographic studies of 2-year-old Picea abies plants show that in summer leaf assimilates from the second-year shoot are translocated basipetally. Leaf assimilates are first transported to the stem via leaf trace phloem, then to the base of the stem in the sieve cells of the latest increment of secondary phloem. On the way down leaf assimilates move radially from sieve cells into cells of the phloem parenchyma, the vascular cambium, the rays, the inner periderm and certain cells of pith and cortex, including the epithelial cells surrounding the resin ducts. Other cells of pith and cortex remain nearly free of label, despite the long translocation time (20 h). With the exception of the vascular cambial cells, the stem cells that gain leaf assimilates by radial distribution coincide with those that contain chlorophyll and starch.     

The Importance of Phellogen Cells and their Structural Characteristics in Susceptibility and Resistance to Excoriation in Immature and Mature Potato Tuber (Solanum tuberosum L.) Periderm

Potato tuber (Solanum tuberosum L.) periderm maturation is animportant physiological process that directly affects the susceptibilityand development of resistance to costly excoriation (skinning-typewounds) at harvest. The objectives of this research were toidentify the specific types of cells and the cellular changesassociated with susceptibility and resistance to tuber excoriationin immature and mature tubers respectively. Epifluorescent microscopicexamination of immature tuber periderm (phellem, phellogen andphelloderm cells) from several genetically diverse cultivarshas shown that the cellular damage resulting from excoriationoccurs within the phellogen (cork cambium), a meristematic layerof cells that gives rise to neighbouring phellem and phellodermcells. Tuber excoriation is the result of the fracture of radialphellogen cell walls linking the skin (phellem) to the phelloderm.As the tuber periderm matures, phellogen cells become inactiveand the radial walls of these cells become more resistant tofracture; resistance to excoriation develops. Ultrastructuralstudies of immature tuber periderm show that radial walls ofactive phellogen cells are thin and fragile. During peridermmaturation, both radial and tangential phellogen cell wallsthicken as they strengthen and become resistant to fracture,thereby providing resistance to excoriation. These results refuteprevious theories of the physiological changes responsible forthe onset of resistance to tuber skinning injury. The combinedresults establish a paradigm whereby the thickening and strengtheningof tuber phellogen cell walls upon periderm maturation are thedeterminant for resistance to tuber excoriation. Copyright 2001Annals of Botany Company Cambium, meristematic, periderm, phellem, phelloderm, phellogen, potato, skinning, Solanum tuberosum L., 0tuber     

油松茎次生结构中树脂道的结构分布和发育的研究

油松茎的次生结构中树脂道存在于次生维管组织中。其中,次生木质部内具有水平的和垂直的两类树脂道,而次生韧皮部内则仅有水平的树脂道。两类树脂道都由上皮细胞和鞘细胞包围着胞间道构成,其中木质部内的树脂道具有死鞘细胞,而韧皮部中的则都系生活细胞。在心材中,垂直树脂道形成拟侵填体。在次生木质部内,垂直树脂道常分布于早材的外部区域和最初形成的晚材中,它们与水平树脂道连接,腔道贯通,从而形成二维网状结构。垂直树脂道来源于纺锤状原始细胞的衍生细胞,而水平树脂道来源于射线原始细胞,两者都以裂生方式发生。