灵武长枣果实维管束韧皮部及周围细胞的超微结构特征

为了探讨灵武长枣果实光合同化物韧皮部卸载和运输的途径,该研究采用透射电镜技术,对不同发育时期灵武长枣果实维管束韧皮部及其周围薄壁细胞的超微结构特征进行了分析。结果表明:筛管/伴胞复合体及其周围韧皮薄壁细胞间在果实膨大前期富含胞间连丝,而韧皮薄壁细胞与周围库细胞以及相邻库细胞间几乎不存在胞间连丝,形成共质体隔离;筛管/伴胞复合体及其与周围薄壁细胞间在果实快速膨大期也存在胞间连丝,但与果实膨大前期相比明显减少;果实着色期,SE/CC复合体及其与周围薄壁细胞间胞间连丝较少,并且出现阻塞现象;果实完熟期,筛管和伴胞之间几乎没有胞间连丝,有的筛管之间有少量胞间连丝,但却出现了阻塞现象,果肉库薄壁细胞与韧皮薄壁细胞间因胞间连丝阻塞现象而形成共质体隔离。综上结果认为,在果实发育的膨大前期阶段,光合同化物以共质体途径经筛分子卸出,卸出后可能以质外体途径进入液泡贮藏与利用;果实快速膨大期,光合同化物的卸出与运输采用共质体和质外体共存的途径;果实着色期和完熟期,光合同化物从筛分子卸出到贮藏薄壁细胞的运输均以质外体途径为主。     

Localization of ATPase in Sink Tissues of Ricinus

Histochemical localization of ATPase was carried out on phloemtissues from vegetative and reproductive sinks of Ricinus communis,using lead precipitation procedures. Reaction products werelocalized mainly at the plasma membrane of the sieve elements,companion cells and phloem parenchyma cells. Activity was alsopresent in plasmodesmata, the tonoplast of companion cells anddispersed P-protein within the sieve element lumen. The resultsare discussed in relation to the possible involvement of a plasmamembrane ATPase in apoplastic and symplastic unloading fromthe phloem conducting tissues. ATPase, sink tissues, unloading, Ricinus communis     

Intercellular Symplastic Connection and Isolation of the Unloading Zone in Flesh of the Developing Grape Berry

The uhrastructure and intercellular connection of the sugar unloading zone (i. e. the phloem in the dorsal vascular bundle and the phloem-surrounding the assimilate sink-cells) of grape ( Vitis vinifera x V. labrusca cv. Jingchao) berry was observed via transmission electron microscopy. The results showed that during the early developmental stages of grape berry, numerous plasmodesmata were found in the phloem between sieve element (SE) and companion cell (CC), between SE/CC complexes, between SE/CC complex and phloem parenchyma cell and in between phloem parenchyma cells, which made the phloem a symplastic integration, facilitating sugar unloading from sieve elements into both companion cells and phloem parenchyma cells via a symplastic pathway. On the contrary, there was almost no plasmodesma between phloem and its surrounding flesh photoassimilate sink-cells, neither in between the flesh photoassimilate sink-cells giving rise to a symplastic isolation both between phloem and its surrounding flesh photoassimilate sink-cells, as well as among the flesh photoassimilate sink-cells. This indicated that both the sugar unloading from phloem and pestphloem transport of sugars should be mainly via an apoplastic pathway. Dining the ripening stage, most of the plasmodesmata between SE/CC complex and the surrounding phloem parenchyma cells were shown to be blocked by the electron-opaque globules, and a phenomenon of plasmolysis was found in a number of companion cells, indicating a symplastic isolation between SE/CC complex and its surrounding parenchynm cells during this phase. The symplastic isolation between the whole phloem and its surrounding photoassimilate sink-cells during the early developmental stages shifted to a symplastic isolation within the phloem during the ripening phase, and thus the symplastic pathway of sugar unloading from SE/CC complex during the early development stages should be replaced by a dominant apoplastic unloading pathway from SE/CC complex in concordance.     

灵武长枣果实ATPase超微细胞化学定位和功能研究

采用ATPase超微细胞化学定位技术,研究灵武长枣果实不同发育阶段韧皮部和果肉库薄壁细胞ATPase分布特征,以明确灵武长枣果实ATPase超微细胞化学定位特征和功能。结果显示:(1)第一次快速生长期SE/CC复合体与周围的薄壁细胞有丰富的胞间连丝,形成共质体连续,韧皮部薄壁细胞之间有丰富的胞间连丝,ATPase反应物在韧皮部各细胞分布较少。(2)缓慢生长期ATPase反应物在韧皮部各细胞分布逐渐增加。(3)第二次快速生长期SE/CC复合体与周围的薄壁细胞缺乏胞间连丝,形成共质体隔离,韧皮薄壁细胞及果肉库薄壁细胞的胞间连丝较少,囊泡和膜泡在筛管、韧皮薄壁细胞和库薄壁细胞中很丰富,质膜、液泡膜、囊泡膜、细胞壁和胞间隙的ATPase活性较高。研究表明,果实在第一次快速生长期同化物从筛分子的卸出主要采取共质体途径,缓慢生长期同化物卸出时可能为共质体和质外体途径共存,第二次快速生长期则主要以质外体途径为主,证明果实不同发育阶段韧皮部同化物卸出路径存在差异。     

What Is Phloem Unloading?

Several studies of phloem unloading have failed to distinguish between transport events occurring at the sieve element/companion cell boundary and subsequent short-distance transport through parenchyma cells. Indirect evidence has been obtained for symplastic unloading in storage and utilization sinks. In other sinks transfer to the apoplast may occur, but not necessarily at the sieve element/companion cell complex, and the evidence for apoplastic phloem unloading is equivocal, as is the role of apoplastic acid invertase in this process. The ability of several types of sink cells to accumulate sugars from the apoplast is discussed in the conflicting light of functional symplastic continuity between sink cells. Attention is drawn to the complexity of the postunloading pathway in many sinks and the difficulty of determining the exact sites of symplast/apoplast solute exchange. Potential future areas for study in the field are highlighted.     

FINE STRUCTURE OF PHLOEM CELLS IN RELATION TO TRANSLOCATION IN THE COTTON SEEDLING

When special precautions were taken to permit killing and fixation of sieve elements before they were cut, sieve pores were found to be open. Companion cells were shown to be highly resistant to freezing injury and less plasmolyzable than phloem parenchyma. Plasmodesmata connected parenchyma to parenchyma, parenchyma to companion cells, and companion cells to sieve elements. Their general absence between parenchyma cells and sieve elements points to a specific role of companion cells in sieve tube functioning. EM studies of these cells revealed an ER system which connects the central core of the plasmodesma to the sieve tube. This system may be responsible for active sucrose transport. Callose was always present on sieve plates of mature functioning sieve elements even with the most rapid killing and fixing possible. Extra callose promoted by heating (45 C) an intact stem segment was found to constrict the sieve pores almost completely. Constriction of plasmodesmata in lateral sieve areas also was evident. Fine structure analysis of the blocking mechanism is in accord with evidence obtained by tracer studies.     

Distribution and frequency of plasmodesmata in relation to photoassimilate pathways and phloem loading in the barley leaf

Large, intermediate, and small bundles and contiguous tissues of the leaf blade of Hordeum tvulgare L. ‘Morex’ were examined with the transmission electron microscope to determine their cellular composition and the distribution and frequency of the plasmodesmata between the various cell combinations. Plasmodesmata are abundant at the mesophyll/parenchymatous bundle sheath, parenchymatous bundle sheath/mestome sheath, and mestome sheath/vascular parenchyma cell interfaces. Within the bundles, plasmodesmata are also abundant between vascular parenchyma cells, which occupy most of the interface between the sieve tube-companion cell complexes and the mestome sheath. Other vascular parenchyma cells commonly separate the thick-walled sieve tubes from the sieve tube-companion cell complexes. Plasmodesmatal frequencies between all remaining cell combinations of the vascular tissues are very low, even between the thin-walled sieve tubes and their associated companion cells. Both the sieve tube-companion cell complexes and the thick-walled sieve tubes, which lack companion cells, are virtually isolated symplastically from the rest of the leaf. Data on plamodesmatal frequency between protophloem sieve tubes and other cell types in intermediate and large bundles indicate that they (and their associated companion cells, when present) are also isolated symplastically from the rest of the leaf. Collectively, these data indicate that both phloem loading and unloading in the barley leaf involve apoplastic mechanisms.     

STUDIES ON THE ROOT OF HORDEUM VULGARE L.– ULTRASTRUCTURE OF THE SEMINAL ROOT WITH SPECIAL REFERENCE TO THE PHLOEM

Seminal root tissue of Hordeum vulgare L. var. Barsoy was fixed in glutaraldehyde and osmium tetroxide and studied with the light and electron microscopes. The roots consist of an epidermis, 6–7 layers of cortical cells, a uniseriate endodermis and a central vascular cylinder. Cytologically, the cortical and endodermal cells are similar except for the presence of tubular-like invaginations of the plasmalemma, especially near the plasmodesmata, in the former. The vascular cylinder consists of a uniseriate pericycle surrounding 6–9 phloem strands occurring on alternating radii with an equal number of xylem bundles. The center of the root contains a single, late maturing metaxylem vessel element. Each phloem strand consists of one protophloem sieve element, two companion cells and 1–3 metaphloem sieve elements. The protophloem element and companion cells are contiguous with the pericycle. Metaphloem sieve elements are contiguous with companion cells and are separated from tracheary elements by xylem parenchyma cells. The protoplasts of contiguous cells of the root are joined by various numbers of cytoplasmic connections. With the exception of the pore-plasmodesmata connections between sieve-tube members and parenchymatic elements, the plasmodesmata between various cell types are similar in structure. The distribution of plasmodesmata supports a symplastic pathway for organic solute unloading and transport from the phloem to the cortex. Based on the arrangement of cell types and plasmodesmatal frequencies between various cell types of the root, the major symplastic pathway from sieve elements to cortex appears to be via the companion and xylem parenchyma cells.     

Symplastic connection is required for bud outgrowth following dormancy in potato (Solanum tuberosum L.) tubers

To gain greater insight into the mechanism of dormancy release in the potato tuber, an investigation into physiological and biochemical changes in tuber and bud tissues during the transition from bud dormancy (immediately after harvest) to active bud growth was undertaken. Within the tuber, a rapid shift from storage metabolism (starch synthesis) to reserve mobilization within days of detachment from the mother plant suggested transition from sink to source. Over the same period, a shift in the pattern of [U-(14)C]sucrose uptake by tuber discs from diffuse to punctate accumulation was consistent with a transition from phloem unloading to phloem loading within the tuber parenchyma. There were no gross differences in metabolic capacity between resting and actively growing tuber buds as determined by [U-(14)C]glucose labelling. However, marked differences in metabolite pools were observed with large increases in starch and sucrose, and the accumulation of several organic acids in growing buds. Carboxyfluorescein labelling of tubers clearly demonstrated strong symplastic connection in actively growing buds and symplastic isolation in resting buds. It is proposed that potato tubers rapidly undergo metabolic transitions consistent with bud outgrowth; however, growth is initially prevented by substrate limitation mediated via symplastic isolation.     

Ultrastructural indications for coexistence of symplastic and apoplastic phloem loading in Commelina benghalensis leaves

The ultrastructural ontogeny of Commelina benghalensis minor-vein elements was followed. The mature minor vein has a restricted number of elements: a sheath of six to eight mestome cells encloses one xylem vessel, three to five vascular parenchyma cells, a companion cell, a thin-walled protophloem sieve-tube member and a thick-walled metaphloem sieve-tube member. The protophloem sieve-tube member (diameter 4–5 m; wall thickness 0.12 m) and the companion cell originated from a common mother cell. The metaphloem sieve-tube member (diameter 3 m; wall thickness 0.2 m) developed from the same precursor cell as the phloem parenchyma cells. Counting the plasmodesmatal frequencies demonstrated a symplastic continuum from mesophyll to the minor-vein phloem. The metaphloem sievetube member and the phloem parenchyma cells are the termini of this symplast. The protophloem sieve-tube member and companion cell constitute an insulated symplastic domain. The symplastic route, mesophyll to metaphloem sieve tube, appears to offer a path for symplastic loading; the protophloem sieve tube may be capable of accumulation from the apoplast. A similar two-way system of loading may exist in a number of plant families. Plasmodesmograms (a novel way to depict cell elements, plasmodesmatal frequencies and vein architecture) of some other species also displayed the anatomical requirements for two routes from mesophyll to sieve tube and indicate the potential coexistence of symplastic and apoplastic loading.     

A morphometric analysis of the phloem-unloading pathway in developing tobacco leaves

A morphometric analysis of developing leaves of Nicotiana tabacum L. was conducted to determine whether imported photoassimilates could be unloaded by symplastic transport and whether interruption of symplastic transport could account for termination of import. Five classes of veins were recognized, based on numbers of cells in transverse section. Photoassimilate is unloaded primarily from Class III veins in tissue nearing the end of the sink phase of development. Smaller veins (Class IV and V) do not transport or unload photoassimilate in sink tissue because the sieve elements of these veins are immature until after the tissue stops importing. In Class III veins the sieve element-companion cell (SE-CC) complexes are surrounded by phloem parenchyma which abuts the bundle sheath. Along the most obvious unloading route, from SE-CC complex to phloem parenchyma to bundle sheath to mesophyll cells, the frequency of plasmodesmata at each interface increases. To determine whether this pattern of plasmodesmatal contact is consistent with symplastic unloading we first demonstrated, by derivation from Fick's law that the rate of diffusion from a compartment is proportional to a number N which is equal to the ratio of surface area to volume of the compartment multiplied by the frequency of pores (plasmodesmata) which connect it to the next compartment. N was calculated for each compartment within the vein which has the SE-CC complex as its center, and was shown to be statistically the same in all cases except one. These observations are consistent with a symplastic unloading route. As the leaf tissue matures and stops importing, plasmodesmatal frequency along the unloading route decreases and contact area between cells also decreases as intercellular spaces enlarge. As a result, the number of plasmodesmata between the SE-CC complex and the first layer of mesophyll cells declines in nonimporting tissue to 34% of the number found in importing tissue, indicating that loss of symplastic continuity between the phloem and surrounding cells plays a role in termination of photoassimilate unloading.Abbreviation SE-CC sieve element-companion cell     

The sucrose transporter StSUT1 localizes to sieve elements in potato tuber phloem and influences tuber physiology and development

The sucrose (Suc) H(+)-cotransporter StSUT1 from potato (Solanum tuberosum), which is essential for long-distance transport of Suc and assumed to play a role in phloem loading in mature leaves, was found to be expressed in sink tubers. To answer the question of whether SUT1 serves a function in phloem unloading in tubers, the promoter was fused to gusA and expression was analyzed in transgenic potato. SUT1 expression was unexpectedly detected not in tuber parenchyma but in the phloem of sink tubers. Immunolocalization demonstrated that StSUT1 protein was present only in sieve elements of sink tubers, cells normally involved in export of Suc from the phloem to supply developing tubers, raising the question of the role of SUT1 in tubers. SUT1 expression was inhibited by antisense in transgenic potato plants using a class I patatin promoter B33, which is primarily expressed in the phloem of developing tubers. Reduced SUT1 expression in tubers did not affect aboveground organs but led to reduced fresh weight accumulation during early stages of tuber development, indicating that in this phase SUT1 plays an important role for sugar transport. Changes in Suc- and starch-modifying enzyme activities and metabolite profiles are consistent with the developmental switch in unloading mechanisms. Altogether, the findings may suggest a role of SUT1 in retrieval of Suc from the apoplasm, thereby regulating the osmotic potential in the extracellular space, or a direct role in phloem unloading acting as a phloem exporter transferring Suc from the sieve elements into the apoplasm.     

Tuberization in potato involves a switch from apoplastic to symplastic phloem unloading

Phloem unloading was studied in potato plants in real time during the early stages of tuberization using carboxyfluorescein (CF) as a phloem-mobile tracer, and the unloading pattern was compared with autoradiography of tubers that had transported (14)C assimilates. In stolons undergoing extension growth, apoplastic phloem unloading predominated. However, during the first visible signs of tuberization, a transition occurred from apoplastic to symplastic transport, and both CF and (14)C assimilates subsequently followed identical patterns of phloem unloading. It is suggested that the switch to symplastic sucrose unloading may be responsible for the upregulation of several genes involved in sucrose metabolism. A detailed analysis of sugar levels and (14)C sugar partitioning in tuberizing stolons revealed a distinct difference between the apical region of the tuber and the subapical region. Analysis of invertase activity in nontuberizing and tuberizing stolons revealed a marked decline in soluble invertase in the subapical region of swelling stolons, consistent with the switch from apoplastic to symplastic unloading. However, cell wall-bound invertase activity remained high in the apical 1 to 2 mm of tuberizing stolons. Histochemical analysis of potato lines transformed with the promoter of an apoplastic invertase gene (invGE) linked to a reporter gene also revealed discrete gene expression in the apical bud region. Evidence is presented that the apical and lateral tuber buds function as isolated domains with respect to sucrose unloading and metabolism.     

宁夏枸杞果实韧皮部及其周围细胞超微结构研究

应用透射电镜技术研究了宁夏枸杞果实韧皮部细胞的超微结构变化。结果表明:(1)随着枸杞果实的发育成熟,果实维管组织中的韧皮部筛分子筛域逐渐变宽,筛孔大而多,通过筛孔的物质运输十分活跃;筛分子和伴胞间有胞间连丝联系,伴胞属传递细胞类型,与其相邻韧皮薄壁细胞和果肉薄壁细胞连接处的细胞界面发生质膜内突,整个筛分子/伴胞复合体与韧皮薄壁细胞之间形成共质体隔离,韧皮部糖分的卸载方式主要以质外体途径进行。(2)韧皮薄壁细胞间的胞间连丝较多,而韧皮薄壁细胞与果肉薄壁细胞的胞间连丝相对较少,但果肉薄壁细胞间几乎无胞间连丝;果肉薄壁细胞之间胞间隙较大,细胞壁和质膜内突间形成较大的质外体空间,为质外体的糖分运输创造了条件。(3)筛管、伴胞、韧皮薄壁细胞和果肉薄壁细胞中丰富的囊泡以及活跃的囊泡运输现象,暗示囊泡也参与了果实糖分的运输过程。研究推测,枸杞果实韧皮部同化物的卸载方式以及卸载后的同化物运输主要以质外体途径为主。     

Minor vein structure and sugar transport in Arabidopsis thaliana

Leaf and minor vein structure were studied in Arabidopsis thaliana (L.) Heynh. to gain insight into the mechanism(s) of phloem loading. Vein density (length of veins per unit leaf area) is extremely low. Almost all veins are intimately associated with the mesophyll and are probably involved in loading. In transverse sections of veins there are, on average, two companion cells for each sieve element. Phloem parenchyma cells appear to be specialized for delivery of photoassimilate from the bundle sheath to sieve element-companion cell complexes: they make numerous contacts with the bundle sheath and with companion cells and they have transfer cell wall ingrowths where they are in contact with sieve elements. Plasmodesmatal frequencies are high at interfaces involving phloem parenchyma cells. The plasmodesmata between phloem parenchyma cells and companion cells are structurally distinct in that there are several branches on the phloem parenchyma cell side of the wall and only one branch on the companion cell side. Most of the translocated sugar in A. thaliana is sucrose, but raffinose is also transported. Based on structural evidence, the most likely route of sucrose transport is from bundle sheath to phloem parenchyma cells through plasmodesmata, followed by efflux into the apoplasm across wall ingrowths and carrier-mediated uptake into the sieve element-companion cell complex. Received: 5 October 1999 / Accepted: 20 November 1999     

Adenosine triphosphatase in the phloem of Cucurbita

Summary The distribution of adenosine triphosphatase (ATPase) activity in the phloem of petioles and minor veins of Cucurbita maxima has been studied using a lead phosphate precipitation procedure. ATPase activity was localized in sieve elements, companion cells and parenchyma cells. Activity was found at the cell surfaces, associated with the dispersed P-protein of mature sieve elements, in mitochondria, sieve-element reticulum, and at specific regions of the cell walls. It is suggested that the ATPase at the phloem cell surfaces may function in intercellular transport of assimilates or ions, and that the ATPase activity associated with the P-protein may function in the translocation process or in callose deposition.     

Histochemical localization of nucleoside triphosphatase activity in assimilate conducting plant tissue

Summary For the histochemical localization of nucleoside triphosphatases at the electron microscopic level, prefixed tissues were incubated with lead nitrate in addition to substrate (GOMORI reaction). While ATP and UTP as substrates gave electron-dense reaction products at the plasmalemma of sieve tubes, companion cells and phloem parenchyma cells, and at plasmodesmata in primary pitfields, AMP gave reaction products only at the tonoplast of parenchyma cells. Since electron-dense deposits also occur in cell walls and vacuoles, energy dispersive X-ray microanalysis was used to distinguish between lead deposits and lead-phosphate deposits. The latter were restricted to the symplast. Among the three plant species used, the leaf bundle phloem ofHordeum distichon showed ATPase activity largely restricted to the phloem cells, except for the thickwalled sieve tubes. Some activity also bordered the chloroplasts of the bundle sheath cells. In the C₄ plantGomphrena globosa, ATPase and UTPase activities appeared to be the greater in phloem parenchyma cells than in sieve tubes. In the phloem of youngMonstera deliciosa roots, ATPase occurred not only at the plasmalemma of sieve tubes, but also around sieve-tube plastids. When compared with AMP as substrate, it appears that nucleoside triphosphates are the natural substrates of the enzyme(s) in the plasmalemma of sieve tubes and phloem parenchyma cells.     

Cytochemical localization of ATPase activity in phloem tissues ofRicinus communis

Summary Standard lead precipitation procedures have been used to examine the localization of ATPase activity in phloem tissues ofRicinus communis. Reaction product was localized on the plasma membrane of the companion cells associated with sieve elements and of parenchyma cells in phloem tissues from the leaf, petiole, stem and root. ATPase activity was also present on the plasma membrane and dispersed P-protein of sieve elements in petiole, stem and root tissue, but was absent from the plasma membrane of these cells in the leaf minor veins. Substitution of-glycerophosphate for ATP produced no change in the localization of reaction product in leaf tissue. These findings are discussed in relation to current theories on the mechanism of sugar transport and phloem loading.     

Plasmodesmatal frequency in relation to short-distance transport and phloem loading in leaves of barley (Hordeum vulgare). Phloem is not loaded directly from the symplast

We investigated the phloem loading pathway in barley, by determining plasmodesmatal frequencies at the electron microscope level for both intermediate and small blade bundles of mature barley leaves. Lucifer yellow was injected intercellularly into bundle sheath, vascular parenchyma, and thin-walled sieve tubes. Passage of this symplastically transported dye was monitored with an epifluorescence microscope under blue light. Low plasmodesmatal frequencies endarch to the bundle sheath cells are relatively low for most interfaces terminating at the thin- and thick-walled sieve tubes within this C₃ species. Lack of connections between vascular parenchyma and sieve tubes, and low frequencies (0.5% plasmodesmata per μm cell wall interface) of connections between vascular parenchyma and companion cells, as well as the very low frequency of pore-plasmodesmatal connections between companion cells and sieve tubes in small bundles (0.2% plasmodesmata per μm cell wall interface), suggest that the companion cell-sieve tube complex is symplastically isolated from other vascular parenchyma cells in small bundles. The degree of cellular connectivity and the potential isolation of the companion cell-sieve tube complex was determined electrophysiologically, using an electrometer coupled to microcapillary electrodes. The less negative cell potential (average –52 mV) from mesophyll to the vascular parenchyma cells contrasted sharply with the more negative potential (–122.5 mV) recorded for the companion cell-thin-walled sieve tube complex. Although intercellular injection of lucifer yellow clearly demonstrated rapid (0.75 μm s^-1) longitudinal and radial transport in the bundle sheath-vascular parenchyma complex, as well as from the bundle sheath through transverse veins to adjacent longitudinal veins, we were neither able to detect nor present unequivocal evidence in support of the symplastic connectivity of the sieve tubes to the vascular parenchyma. Injection of the companion cell-sieve tube complex, did not demonstrate backward connectivity to the bundle sheath. We conclude that the low plasmodesmatal frequencies, coupled with a two-domain electropotential zonation configuration, and the negative transport experiments using lucifer yellow, precludes symplastic phloem loading in barley leaves.     

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