The Ultrastructure of the Digestive Glands in Pinguicula vulgaris L. (Lentibulariaceae) Relative to their Function. II. The Changes on Stimulation

The secretory cells of the digestive glands remain highly activeduring the entire period of prey digestion and absorption ofnutrients. They appear to play a major role in gland activity.A model of the digestive gland's activity on stimulation isproposed. It is very similar to that suggested earlier for Dionaeamuscipula. After the digestion and absorption cycle, destructiveprocesses are initiated in the glands. These appear similarto those observed in the glands of the ageing, unstimulatedleaf and are not associated with feeding. Pinguicula vulgaris L. carnivorous plant, digestive glands, ultrastructure, protein secretion absorption, senescence     

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     

The Ultrastructure of Ion-Secreting and Non-Secreting Salt Glands of Limonium platyphyllum

The ultrastructure of salt glands in developing leaves of Limoniumplatyphyllum is described prior to exposure to 3% NaCl solution(with non-secreting glands) and after 4.5 and 18 h exposureto the salt solution. It is shown that in most glands, the transitionto active chloride transport was accompanied by the displacementof vacuoles toward the cell periphery and by the establishmentof plasmalemma contact sites with the tonoplast which appearedsimilar to gap junctions in animal epithelial cells. No evidencefor the exocytosis of vacuoles was found. It is suggested thatgland vacuoles may have a primary role in chloride secretionand that the tonoplast may be functionally asymmetrical, sothat the free part facing the hyaloplasm bears ion pumps, whereashighly permeable ion channels are active along the zone of contactwith the plasmalemma. It follows that the active step in chloridetransport in Limonium glands is the influx of ions into thevacuoles. Within the inner cup cells of the gland, vacuolescome into contact with the plasmalemma only at sites where thecell wall is adjacent to secretory and accessory cells. Suchan asymmetry appears to ensure the directed flux of ions intothis cell wall. Wall protuberances in the gland cells are rudimentaryand presumably not involved directly in NaCl secretion. Thenucleolus is activated during secretion and the frequency offree ribosomes is significantly increased, which is suggestiveof their involvement in the synthesis of membrane transportproteins. The ultrastructure of about one-third of the glandsremained unchanged in salt-treated leaves. Key words: Salt glands, ultrastructure, ion fluxes     

Ultrastructural features of the salt gland of Tamarix aphylla L.

Summary The salt gland in Tamarix is a complex of eight cells composed of two inner, vacuolate, collecting cells and six outer, densely cytoplasmic, secretory cells. The secretory cells are completely enclosed by a cuticular layer except along part of the walls between the collecting cells and the inner secretory cell. This non-cuticularized wall region is termed the transfusion are (Ruhland, 1915) and numerous plasmodesmata connect the inner secretory cells with the collecting cells in this area. Plasmodesmata also connect the collecting cells with the adjacent mesophyll cells.There are numerous mitochondria in the secretory cells and in different glands they show wide variation in form. In some glands wall protuberances extend into the secretory cells forming a labyrinth-like structure; however, in other glands the protuberances are not extensively developed. Numerous small vacuoles are found in some glands and these generally are distributed around the periphery of the secretory cells in association with the wall protuberances. Further, an unusual structure or interfacial apparatus is located along the anticlinal walls of the inner secretory cells. The general structure of the gland including the cuticular encasement, connecting plasmodesmata, interfacial apparatus, and variations in mitochondria, vacuoles, and wall structures are discussed in relation to general glandular function.     

Ontogenesis of Tannin Coenocytes in Sambucus racemosa L. II. Mother Tannin Cells

Tannin coenocytes develop from mononucleate tannin mother cells.The process occurs within the whole of the first (youngest)internode and its development can be divided into three stages.In stage I the MTC is isodiametric and similar to the surroundingcells of the flank meristem, being present in the ninth celllayer from the apex surface. The nucleus becomes lanceolateand elongates, and large cytoplasmic vacuoles appear. A twofoldelongation of both the cell and nucleus continues in the secondstage, the cell-nucleus ratio indicating that it is due to theenlarged vacuole, which pushes a thin layer of cytoplasm closeto the cell wall. In this layer of cytoplasm dilated ER cisternaoccur together with small and large vacuoles, a fusion of thevacuoles increasing their volume. Such cells are diploid inspite of larger nuclear volume and rough structure of its chromatin. Sambucus racemosa L., tannin cells, development, ontogenesis     

Pathways for Nutrient Transport in the Pitchers of the Carnivorous Plant Nepenthes alata

Glands of the carnivorous pitcher plant Nepenthesalata are activein transport of materials into and out of the pitcher lumen,indicating dual functions in both secretion and absorption.This study examined the potential for open transport throughthese glands using the ultrastructural tracer lanthanum, whichis restricted to the apoplast, and the fluorescent symplastictracer, 6(5)carboxyfluorescein. Glandular uptake of lanthanumfrom the pitcher fluid occurred through the outer cell wallbetween irregularly spaced cutinized deposits, but was blockedfrom entering the underlying mesophyll cell walls by thick endodermal-likeregions. Similarly, lanthanum localization showed an open apoplasticpathway from the petiole to the endodermal regions in the glandbase. Thus, transport of materials into or out of the glandmust occur through the symplast. 6(5)Carboxyfluorescein showedthat these glands transport fluids directly from the pitcherfluid into vascular endings immediately beneath them via a symplasticroute. When applied to the petiolar vascular system, the fluorescenttracer freely entered immature pitchers, but was blocked fromentering the lumen of the mature pitcher by an endodermal zone.An ultrastructural survey showed infrequent pits with plasmodesmatalconnections to adjoining subepidermal cells. These results indicatethat the function of the gland is developmentally regulated.Prior to maturity, the primary function of the gland appearsto be secretion. However, at maturity, secretion is blockedby an endodermal layer, which limits the function of the glandto absorption. These studies support the theory that the glandsof Nepenthesalata are specialized for the bi-directional transportof materials.Copyright 1999 Annals of Botany Company Apoplastic transport, 6(5)carboxyfluorescein, carnivorous plants, digestive glands, endodermal layer,Nepenthesalata Blanco, lanthanum, pitcher plants.     

Cellular pathway of photosynthate transport in coats of developing seed of Vicia faba L. and Phaseolus vulgaris L. II. Principal cellular site(s) of efflux

The cells responsible for the photosynthate efflux from coatsof developing seed of Vicia faba L. and Phaseolus vulgaris L.were elucidated using known properties of the efflux mechanism.Sensitivity of sucrose efflux to NEM and high potassium concentrationswas retained by seed-coat halves of Phaseolus following pectinaseremoval of the branch parenchyma cell layer. In contrast, removalof the thin-walled parenchyma transfer cell layer from Viciaseed-coat halves abolished this sensitivity. The membrane-impermeantthiol-binding fluorochrome, qBBr, selectively stained the surfaceof the thin-walled parenchyma transfer cells. This phenomenonwas inhibited by the slowly permeable sul-phydryl agent, PCMBS,indicating that the plasma membranes of these cells are enrichedin sulphydryl groups characteristic of membrance porter proteins.On the basis that carrier-mediated sucrose efflux from seedcoats appears to be proton coupled, the putative plasma membraneH+-ATPase was used as a marker for the cells responsible forcarrier-mediated photosynthate efflux. When seed-coat halveswere exposed briefly at pH 8.5 to the weak acid fluorochrome,SRG, the ground parenchyma and thin-walled parenchyma transfercell layers selectively accumulated the dye. The apparent lowpH environment in the walls of these cells that renders SRGmembrane permeant appeared to be maintained by a VAN-sensitiveproton pump. The findings with SRG were corroborated by thecyto-chemical localization of plasma membrane ATPase activityto the ground parenchyma and thin-walled parenchyma transfercells using precipitation of cerium phosphate. Together, ourobservations provide qualified support for the conclusion thatcarrier-mediated photosynthate efflux from coats of Phaseolusand Vicia seed is primarily restricted to the ground parenchymaand thin-walled parenchyma transfer cell layers, respectively. Key words: Ground parenchyma, Phaseolus vulgaris L., photosynthate efflux, seed coat, transfer cell, Vicia faba L.     

Fine Structure and Cytochemistry of the Abscission Zone Cells of Phaseolus Leaves: II. Localization of Peroxidase and Acid Phosphatase in the Separation Zone Cells

The sub-cellular localization of acid phosphatase and peroxidaseactivities has been studied by cytochemical procedures in cellsat the surface of the separation layer during the abscissionof leaves of Phaseolus vulgaris. Intense staining for both enzymeswas found in the cell walls, Golgi bodies and endoplasmic reticulum.The wall staining for acid phosphatase was chiefly associatedwith the middle lamellar region while staining for peroxidasewas found throughout the wall. These observations are discussedin relation to the possible involvement of these enzymes inthe changes occurring in the wall during abscission and to therole of the Golgi bodies in the separation process.     

Pathway of Photosynthate Transfer in the Developing Seed of Vicia faba L: A Structural Assessment of the Role of Transfer Cells in Unloading from the Seed Coat

Photosynthate movement within the coat of the developing seedof Vicia faba occurs radially inward from the restricted vascularsystem and laterally through the non-vascularized region ofthe seed coat prior to exchange to the seed apoplast. Thin-walledparenchyma/transfer cells line the entire inner surface of theseed coat and thus are located at the terminus of the photosynthatetransfer pathway. The principal cellular route of transfer withinthe seed coat and the role of the thin-walled parenchyma/transfercells in membrane exchange to the seed apoplast has been investigated.Sucrose fluxes, computed from estimates of the plasma membranesurface areas of the cell types of the pathway, the plasmodesmatalcross-sectional areas interconnecting contiguous cells and theobserved rate of sucrose delivery to the embryo indicate thatsieve element unloading and subsequent transfer to the thin-walledparenchyma/transfer cells is through the symplast. For the cellsof the ground tissue, plasmodesmatal density is consistentlyhigher on their anticlinal walls. This observation supportsthe reported pattern of lateral transfer through these tissuesin the non-vascular regions of the seed coat. Wall ingrowthsare initiated sequentially in the thin-walled parenchyma cellsto maintain 1–3 rows of thin-walled parenchyma/transfercells. The development of these wall ingrowths results in a58% increase in the plasma membrane surface area of these cellsand provides them with the capacity to act as the principalcellular site for membrane exchange of sucrose to the seed apoplast.This cellular route of symplastic transfer from the sieve elementsto the ground tissues where membrane exchange to the seed apoplastoccurs is consistent with that reported for Phaseolus vulgaris Key words: Cellular pathway, photosynthate transfer, transfer cell, Vicia seed coat     

Comparative studies onChlorella cell walls: Induction of protoplast formation

Among 12 strains ofChlorella ellipsoidea, C. vulgaris, andC. saccharophila tested, 4 strains (1,C. ellpsoidea; 2,C. vulgaris; 1,C. saccharophila) formed osmotically labile protoplasts after treatment with mixtures of polysaccharide degrading enzymes. The relationship between enzymatical digestibility and structure or composition ofChlorella cell walls were studied by electron microscopy and staining techniques with some specific dyes. The cell wall structures of the 12Chlorella strains were grouped into three types: (1) with a trilaminar outer layer, (2) with a thin outer monolayer, and (3) without an outer layer. Protoplasts were formed only from the strains with a cell wall of Type 2. In the strains with a cell wall of Type 1, the outer layer protected the inner major microfibrillar layer against enzymatic digestion. The cell wall of Type 3 was totally resistant to the enzymes; the chemical composition of the cell wall would be somewhat different from that of other types.     

Response to Phosphate Deficiency in Bean (Phaseolus vulgarisL.) Roots. Respiratory Metabolism, Sugar Localization and Changes in Ultrastructure of Bean Root Cells

Bean plants (Phaseolus vulgarisL. ‘Zlota Saxa’)were cultured on complete (+P) or phosphate-deficient (-P) nutrientmedium. A large increase in glucose concentration was foundin the meristematic zone of -P roots compared to control roots.The increased glucose concentration in the meristematic zonedid not influence total respiration rate. Glucose or uncoupler(carbonyl cyanide m-chlorophenylhydrazone) failed to increasethe respiration rate in -P root segments, but stimulated respirationin +P roots. The ultrastructure of cortical cells from the meristematicroot zone showed marked differences between +P and -P plants.Large vacuoles, invaginations of the plasmalemma and condensedforms of mitochondria were dominating features in cortical cellsof -P roots. Analysis of extracts after treating roots withdimethylsulfoxide (DMSO) indicated different localization ofsugars in the cell compartments. In roots of -P plants, mostof the reducing sugars were detected in the cytoplasm fractionwhile most sucrose was in the vacuole. Observations of the effectof 10% DMSO on cell ultrastructure indicated partial destructionof the plasmalemma but not the tonoplast. The localization ofreducing sugars in secondary vacuoles or plasmalemma invaginationsin the cells from the meristematic region of -P roots is discussed.Copyright1998 Annals of Botany Company. Bean (Phaseolus vulgarisL.), roots, Pi deficiency, respiration, meristematic zone, ultrastructure, sugar efflux, reducing sugars and sucrose localization.     

Isolation and characterization of the cell wall receptor of 14C-malformin in Phaseolus vulgaris L.

¹⁴C-malformin attaches to at least two cell wall receptors inPhaseolus vulgaris. One receptor was extracted with Tris buffer(pH 8.5) and the other with 0.1 N NaOH. In both cases, priortreatment of the walls with wall degrading enzymes (macerase,cellulysin) was required. The two receptors differed with regardto ultrafiltration and gel filtration chromatography. The Tris-extractedreceptor is a protein, probably a glycoprotein, which containshydroxyproline and sulfhydryl groups. Although cuttings nottreated with malformin had Tris-extractable receptor, formationof the receptor appeared to be enhanced by malformin. ¹ Present address: American Cyanamid, P. O. Box 400, Princeton,New Jersey 08540, U. S. A. (Received August 2, 1976; )     

Cell Wall Degrading Enzymes in Cuscuta reflexa and Its Hosts

Pectin degrading enzymes, hemicellulose degrading enzyme andcellulose degrading enzymes were studied in Cuscuta reflexaRoxb., its susceptible hosts, Brassica campestris L., Cocciniaindica W. & A. Datura innoxia Mill, Helianthus annuus L.,Holoptelea indica Planch, Lantana camara L., Medicago sativaL., Manihot utilissima Pohl, Petunia hybrida X Hort exvilm,Pisum sativum L., Phaseolus vulgaris L. and Solanum nigrum L.and non-susceptible plants Ipomoea batata Lam. and Solanum tuberosumL. Pectin esterase and polygalacturonase were present in higheramounts in Cuscuta parasitic on P. vulgaris and S. nigrum, whichneeded more time for haustorial establishment. Exo-l, 4-ß-D-glucosidaseactivity was found in Cuscuta but could not be detected in itshosts. Xylanase and cellulase activity of host plants increasedwhile cellobiase activity decreased as a result of infectionby the parasite. Higher pectin esterase, polygalacturonase,xylanase and exo-l, 4-ß-D-glucosidase activities inthe haustorial region of the parasite is likely to bring aboutthe lysis of the cell wall of the host plant and thus facilitatethe penetration of the parasite haustoria into the host sieveelement, which is necessary for the transport of nutrients betweenthe host and the parasite. Key words: Cell wall degrading enzymes, Cuscuta reflexa     

Ultrastructure and Development of Oil Idioblasts in Annona muricata L.

The ultrastructure and development of oil idioblasts in theshoot apex and leaves in Annona muricata L. are described, andthree arbitrary developmental stages are distinguished: cellsin which no additional cell wall layers have been depositedagainst the initial primary cell wall, possessing an electron-translucentcytoplasm and distinct plastids which lack thylakoids (stage1); cells in which a suberized layer has been deposited againstthe primary wall (stage 2, the cytoplasm resembles that of thepreceding stage), and cells in which an additional inner walllayer has been deposited against the suberized layer, whichincreases in thickness with development (stage 3). In this stagean oil cavity is formed, surrounded by the plasmalemma, andattached to a bell-like protrusion of the inner wall layer,the cupule. A complex membranous structure occurs next to thecupule. Smooth tubular endoplasmic reticulum (ER), appearingas linearly arranged tubules, and groups of crystalline bodieswith an almost hexagonal outline are present. The final stagewas further subdivided into three subgroups (a, b, c) basedon the extent of the oil cavity, its contents, and the compositionof the cytoplasm, and increasing thickness of the inner walllayer. The oil is probably synthesized in the plastids, releasedinto the cytoplasm, and then passed through the plasmalemmasurrounding the oil cavity. Oil idioblasts, Annona muricata L., suberized layer, inner wall layer, oil cavity, cupule, smooth tubular ER, crystalline bodies     

The Cellular Pathway of Short-distance Transfer of Photosynthates and Potassium in the Elongating Stem of Phaseolus vulgaris L. A Structural Assessment

The potential cellular pathway of radial transfer of photosynthateand potassium delivered in the phloem to the elongation zone(apical 0.5–2.5 cm) of internode 2 ofPhaseolus vulgarisL. seedlings was elucidated. This was achieved using ultrastructuralobservations of the cell types that constitute the radial pathwayand estimates of potential sucrose and potassium fluxes throughthe cross-sectional area of interconnecting plasmodesmata andacross the plasma membrane surface areas of selected cell types.The investigation relied on predicting the relative roles ofthe mature and developing sieve elements as conduits for theaxial delivery of solutes to the elongation zone. In turn, thesepredictions led to formulation of two transport models whichwere subsequently evaluated. It was found that unloading ofsucrose and potassium from the protophloem sieve elements cannotbe through the symplast due to the absence of plasmodesmata.On the other hand, mature metaphloem sieve element-companioncell complexes have the potential capacity to unload eitherthrough the stem symplast or apoplast. The potential symplasticroute is proposed to be via the companion cells to the adjacentlarge phloem parenchyma cells. Continued radial transfer couldoccur either by exchange to the stem apoplast from the largephloem parenchyma cells or continue in the symplast to the groundtissues. It was further predicted that sucrose utilized forthe development of the procambial/small phloem parenchyma cellscould be delivered axially by them and not by the mature sieveelements. Phaseolus vulgaris ; apoplast; elongating stem; photosynthates; potassium; transport; symplast     

A cytochemical study of the leaf-gland enzymes of insectivorous plants of the genus Pinguicula

Summary Cytochemical methods have been used to study the distribution of acid phosphatase, esterase, ribonuclease, amylase and protease activity in the stimulated and unstimulated leaf glands of Pinguicula grandiflora, P. vulgaris, P. lusitanica, and P. caudata. Two gland types are present, stalked and sessile. The stalked glands bear a muco-polysaccharide secretion droplet, and are concerned with capture of the prey; the sessile glands are specialised for digestion. In unstimulated glands of both classes, acid phosphatase, esterase and ribonuclease activity is associated with the anticlinal walls of the head cells, which have a characteristic spongy inner surface, comparable with that of transfer cells. Acid phosphatase and esterase activity was also detected in the vacuoles of the head cells of the sessile glands. Substrate film tests showed that amylase is readily released from the stalked glands but not from the sessile ones, while in contrast proteolytic activity is mainly associated with the sessile glands.On stimulation by suitable nitrogenous materials, the glands begin to sectete fluid onto the leaf surface within 1 hr. During the process the enzymes held in the spongy walls are discharged, and activity is also lost from the intracellular sites in the sessile glands.Digestion on the leaf surface and resorption of the products has been followed autoradiographically after feeding of ¹⁴C-labelled protein. Within 2 hr, digestion products enter the leaf, and move towards the margin in the vascular system. Movement out of the leaf begins within 12 hr. Microautoradiographs showed a concentration of products around the bases of the sessile glands and in the cells of the gland head, showing that these glands are involved in resorption as well as secretion.A possible mechanism of gland function is discussed.     

The Pathway of Radial Transfer of Photosynthate in Decapitated Stems of Phaseolus vulgaris L.

[¹⁴C]Sucrose was found to be the predominant component of the¹⁴C-photosynthates that accumulated in the free space of decapitatedstems of P. vulgaris plants. The ¹⁴C-photosynthates appearedto occupy the entire free-space volume of the stems at totalsugar concentrations in the range of 3–12 mM. The free-spacesugar levels were found to rapidly decline once photosynthatetransfer to the stems was halted. Moreover, it was found thatestimates of the rate of in vitro sucrose uptake by the stemscould account fully for the decline in free-space sugar levels.Overall, the evidence indicated that at least part of the radialpathway of photosynthate transfer in bean stems involved thestem apoplast. It is tentatively proposed that, based on celland tissue distribution of ¹⁴C-photosynthates, the apoplasticpathway extends from the membrane boundary of the sieve element/companion-cellcomplex to all other cells of the stem. Apoplast, Phaseolus vulgaris L., bean, phloem unloading, photosynthates, symplast     

The Cellular Pathway of Short-distance Transfer of Photosynthates and Potassium in the Elongating Stem ofPhaseolus vulgarisL. A Physiological Assessment

Plasmolytic disruption of plasmodesmata interconnecting metaphloemsieve element-companion cell complexes with small and largephloem parenchyma cells in the elongating region of internode2 ofPhaseolus vulgarisL. seedlings did not affect accumulationof phloem-imported¹⁴C-photosynthates and⁸⁶rubidium. The membrane-impermeantdye, 5(6) carboxyfluorescein, loaded into leaf phloem as themembrane-permeant diacetate ester, was found not to move radiallyout of the importing sieve elements in the internode elongationregion. In contrast, the apoplasmic tracer, Calcuofluor White,rapidly moved laterally throughout all tissues of the elongationzone. Hexoses, sucrose and potassium were identified as themain osmotica in internode apoplasmic sap. Label asymmetry in[¹⁴C](fructosyl)sucrose was retained on accumulation by excisedstem segments. Uptake of [¹⁴C]sucrose and⁸⁶rubidium by stemsegments exhibited saturation kinetics. Sucrose uptake was inhibitedby the slowly penetrating sulphydryl reagent, para-chloromercuribenzenesulphonicacid.In vitrorates of sucrose uptake, at apoplasmic concentrations,corresponded to its predictedin vivorate of delivery to thestem ground tissues from mature sieve elements when respiratorylosses were assumed to be confined to the stem phloem. For potassium,the total delivery rate could be accounted for by itsin vitrorateof uptake. Overall, it was concluded that radial transport,in the elongation zone of internode 2 ofPhaseolus vulgarisL.seedlings, follows an apoplasmic route from mature sieve elementsto stem ground tissues.Copyright 1998 Annals of Botany Company PhaseoluLes vulgaris, apoplasm, elongating stem, French bean, photosynthates, potassium, radial transfer, symplasm.     

Cellular Structures, Plasma Membrane Surface Areas and Plasmodesmatal Frequencies of the Stem of Phaseolus vulgaris L. in Relation to Radial Photosynthate Transfer

At an early stage of secondary development, the metaphloem sieveelements appeared to be the only functional axial transportconduit in fully elongated stems of P. vulgaris plants. Thereis no apparent barrier to the radial transfer of solutes inthe stem apoplast. However, radial transfer through the stemsymplast could be limited by discontinuities resulting fromprotoplast degeneration of the protophloem fibres and developingsecondary xylem fibres. Estimates of possible sucrose fluxesthrough the apoplastic and symplastic routes indicated thatradial photosynthate transfer from the sieve element-companioncell (se-cc) complexes of the stem metaphloem could follow eithercellular route. In the case of apoplastic transfer, the plasmamembrane surface area of the se-cc complexes is only sufficientto support some form of facilitated movement of sucrose. Incontrast, the plasma membrane surface area of the phloem parenchymais sufficient to permit passive diffusion of sucrose to theapoplast. Plasmodesmatal frequencies suggest that any symplastictransfer to the phloem parenchyma from the sieve elements wouldbe via the companion cells. Phaseolus vulgaris, french bean, stem, photosynthate, radial transfer (photosynthates), cellular pathway     

Differentiation of the Tapetum During Microsporogenesis in Tomato (Lycopersicon esculentum Mill.), with Special Reference to the Tapetal Cell Wall

During microsporogenesis and pollen maturation, the tapetumin anthers of tomato (Lycopersicon esculentum) underwent severalultrastructural changes and ultimately degenerated. The changesobserved related to the secretory function of the tapetum andto the transfer of materials from the cytoplasm to the surfaceof tapetal cells. Electron dense deposits, initially in thevacuoles, disappeared coincident with the appearance of orbiculeson the cell wall. The fibrillar wall of the tapetal cells loosened,presumably to facilitate transfer of materials through the wall.In Addition, membranous fragments were a consistent featurein the tapetum wall and may play a role in transport of materials.The cells of the inner tapetum (towards the connective) andouter tapetum (towards the epidermis) had different ultrastructuralfeatures. The cytoplasm of the outer tapetum was more electrondense and had a higher proportion of dictyosomes and mitochondriathan the inner tapetum, indicating the greater secretory natureof the outer tapetum. The plastids and mitochondria also differedin morphology between the two regions. Degenerations of thetapetal cytoplasm began by the vacuolate microspore stage. Atanthesis, cytoplasm was absent but the orbicular wall of thetapetum remained appressed to the wall of the middle layer ofthe anther.Copyright 1993, 1999 Academic Press Lycopersicon esculentum, microsporogenesis, pollen development, tapetum development, tomato, ultrastructure