TRANSFER CELLS IN THE PLACENTAL PAD AND CARYOPSIS COAT OF PAPPOPHORUM SUBBULBOSUM ARECH. (POACEAE)

During caryopsis development the layers of the pericarp, integuments, and nucellus all contribute to the formation of the caryopsis coat. The coat consists of a layer of outer pericarp epidermal transfer cells, a collapsed and senescent layer of middle pericarp cells, and a discontinuous layer of inner pericarp epidermal transfer cells. The latter is not present across the placental pad. The integuments are present as a collapsed dense layer, the nucellus is discontinuous and cellular. The placental pad occurs at the ventral surface of the caryopsis, opposite the scutellum and coleorhiza. It consists of 15–20 collapsed cell layers, including the pigment strand and placental vascular bundle. From the inside several partially collapsed cell layers of the nucellar projection occur which contain transfer-cell walls. The middle dense layers, the pigment strand, consist of the middle pericarp remnant, plus the remains of the placental vascular bundle. The pericarp inner epidermis does not extend across the pad. The aleurone layer is a continuous uniseriate layer around the entire caryopsis except at the placental pad; here it is crushed and contains the remnant of a transfer-cell wall. The outer pericarp epidermis is a continuous layer of transfer cells across the pad. These cells contain membranous inclusions suggesting that they may be functional during germination.     

Sink anatomy in relation to solute movement in rice (Oryza sativa L.): A summary of findings

The results of studies on assimilate and water transport in the developing caryopsis of rice are summarised. Evidence is presented for a symplastic movement of solutes as far as the aleurone layer. However, transport into the apoplast at the nucellus/aleurone interface appears to be a necessary step due to the absence of plasmodesmata at this site. It is suggested that water leaves the caryopsis during grain filling by the isolated cell walls of the pigment strand, the suberised walls of these cells functioning to isolate the apoplast from the symplast and thereby allowing opposing fluxes of water and assimilates to occur in the dorsal region of the grain.     

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.     

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。     

Phloem unloading in the potato tuber. Pathways and sites of ATPase

Summary Potential pathways for sucrose unloading in the potato tuber were examined by light and electron microscopy. Abundant plasmodesmata connected sieve elements with surrounding parenchyma elements and also sieve elements with companion cells. Plasmodesmata were rarer, however, between companion cells and parenchyma elements. These observations suggest that sucrose may leave the sieve elements and enter the storage parenchyma cells directly via the symplast and that transport through the companion cell may not be a prerequisite for unloading. Plasmodesmata, grouped together in primary pit fields, were also abundant between storage cells, and isolated storage cells, separated enzymically, showed considerable variation in plasmodesmatal distribution between cells and also on different faces of a single cell. Deposition of starch was found to occur in the tuber cortex while an endodermis with Casparian strip was present external to the phloem, suggesting that assimilates initially enter the cortical storage cells by an entirely symplastic pathway. The possible involvement of ATPase in the unloading process was examined cytochemically, using a lead-salt precipitation method. By contrast with previous findings for phloem no evidence was found for ATPase activity that was unique to the sieve element-companion cell complex. The present observations favour the view that phloem unloading in the potato tuber is a symplastic and passive process.     

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.     

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.     

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。     

The cellular pathway of photosynthate transfer in the developing wheat grain. II. A structural analysis and histochemical studies of the pathway from the crease phloem to the endosperm cavity

In the developing wheat grain, photosynthate is transferred longitudinally along the crease phloem and then laterally into the endosperm cavity through the crease vascular parenchyma, pigment strand and nucellar projection. In order to clarify this cellular pathway of photosynthate unloading, and hence the controlling mechanism of grain filling, the potential for symplastic and apoplastic transfer was examined through structural and histochemical studies on these tissue types. It was found that cells in the crease region from the phloem to the nucellar projection are interconnected by numerous plasmodesmata and have dense cytoplasm with abundant mitochondria. Histochemical studies confirmed that, at the stage of grain development studied, an apoplastic barrier exists in the cell walls of the pigment strand. This barrier is composed of lignin, phenolics and suberin. The potential capacity for symplastic transfer, determined by measuring plasmodesmatal frequencies and computing potential sucrose fluxes through these plasmodesmata, indicated that there is sufficient plasmodesmatal cross-sectional area to support symplastic unloading of photosynthate at the rate required for normal grain growth. The potential capacity for membrane transport of sucrose to the apoplast was assessed by measuring plasma membrane surface areas of the various cell types and computing potential plasma membrane fluxes of sucrose. These fluxes indicated that the combined plasma membrane surface areas of the sieve element–companion cell (se–cc) complexes, vascular parenchyma and pigment strand are not sufficient to allow sucrose transfer to the apoplast at the observed rates. In contrast, the wall ingrowths of the transfer cells in the nucellar projection amplify the membrane surface area up to 22-fold, supporting the observed rates of sucrose transfer into the endosperm cavity. We conclude that photosynthate moves via the symplast from the se–cc complexes to the nucellar projection transfer cells, from where it is transferred across the plasma membrane into the endosperm cavity. The apoplastic barrier in the pigment strand is considered to restrict solute movement to the symplast and block apoplastic solute exchange between maternal and embryonic tissues. The implications of this cellular pathway in relation to the control of photosynthate transfer in the developing grain are discussed.     

Plasmodesmatal distribution and frequency in vascular bundles and contiguous tissues of the leaf ofThemeda triandra

Small and intermediate vascular bundles and contiguous tissues of the leaf blade ofThemeda triandra var.imberbis (Retz.) A. Camus were examined with transmission and scanning electron microscopes to determine the distribution and frequency of plasmodesmata between various cell types. Plasmodesmata are most abundant at the mesophyll/bundle-sheath cell and bundle-sheath/vascular parenchyma cell interfaces, and their numbers decrease with increasing proximity to both thick- and thin-walled sieve tubes. Among cells of the vascular bundles, the greatest frequency of plasmodesmata occurs between vascular parenchyma cells, followed by that of plasmodesmata between vascular parenchyma cells and companion cells, and then by the pore-plasmodesmata connections between companion cells and thin-walled sieve tubes (sieve tube-companion cell complexes). The sieve tube-companion cell complexes of theT. triandra leaf are not isolated symplastically from the rest of the leaf and, in this respect, differ from their counterparts in theZea mays leaf. However, the thick-walled sieve tubes, like their counterparts inZea mays, lack companion cells and are symplastically connected with vascular parenchyma cells that about the xylem.Abbreviations SEM scanning electron microscope - TEM transmission electron microscope     

Vascular Anatomy of Angiospermous Leaves,with Special Consideration of the Maize Leaf

In this brief review an attempt has been made to discuss some of the important features of the vascular anatomy of angiospermous leaves, especially those related to assimilate transport. Accordingly, emphasis has been placed on the small or minor veins, which are closely related spatially to the mesophyll, and which play a major role in the uptake and subsequent transport of photosynthates from the leaf. The small veins are enclosed by bundle sheaths that intervene between the mesophyll and vascular tissues and greatly increase the area for contact with mesophyll cells. In the minor veins of dicotyledonous leaves, parenchymatic cells having organelle-rich protoplasts and numerous cytoplasmic connections with sieve elements dominate quantitatively. It is these so-called intermediary cells that apparently are directly involved with the loading of assimilates into the sieve elements. In the maize leaf the small and intermediate bundles have two types of sieve tubes, relatively thin-walled ones that have numerous cytoplasmic connections with companion cells, and thick-walled ones that lack companion cells but have numerous connections with vascular parenchyma cells. The companion cell-sieve tube complexes are virtually isolated symplastically from other cells of the vascular bundle and from the bundle sheath. Thick-walled sieve tubes similar to those in the maize leaf have been recorded in the leaves of other grasses.     

Vascular Bundle Distribution Within Broad Bean Seed Coat and the Structure of Unloading Region

Cytological observations were made on developing seed coat of broad bean (Vicia faba L.) by use of light and electron microscopy. Attention was focused on vascular distribution. The seeds were attached by the funiculus to tile large vascular bundles of pericarp of broad bean. The vascular bundle passed through hilum and two layers of palisade, entered the pa- renchyma of seed coat, then diverged in to two routes. One was a complete vascular bundle composed of both'phloem and xylem elements, it stretched down through seed raphe, then upward and terminated near the radical. The other was a two-recurrent-vascular-bundle with only phloem constitutents, they extended forward detoured the micropyle and extended downward, but did not join with the upward complete vascular bundle. The recurrent vascular bundles branched out many small short branches. The obvious difference between phloem of recurrent vascular bundle and of complete vascular bundle was that the companion ceils of the former did not normally modify to transfer ceils, but connected to the adjoining parenchyma cells through abundant plasmodesmata. It is deduced from the structural analysis that the symplastic route may play an important role, particularly in the region of recurrent vascular bundle, in the course of importing assimilates unloading in seed coat and transporting to the embryo.     

Transport of assimilates in the developing caryopsis of rice (Oryza sativa L.)

The movement of water in the dorsal region of the developing rice caryopsis was studied using solutions of the heavy metals lanthanum and uranium. In the electron microscope electron-opaque deposits were confined to the cell walls of the pigment strand indicating that this is the main route for the water which enters and leaves the caryopsis during grain filling. The pathway of assimilates into the developing caryopsis was examined using isolated caryopses which had taken up solutions of fluorescent dyes and also by autoradiography of caryopses which had transported ¹⁴C-labelled assimilates in vivo. The results show that assimilates unloaded from the phloem move through the pigment strand and circumferentially via cells of the nucellus before entering the endosperm. A scheme is presented for the interrelations of water and assimilate transport during grain filling     

Leaf structure in relation to solute transport and phloem loading in Zea mays L.

Small and intermediate (longitudinal) vascular bundles of the Zea mays leaf are surrounded by chlorenchymatous bundle sheaths and consist of one or two vessels, variable numbers of vascular parenchyma cells, and two or more sieve tubes some of which are associated with companion cells. Sieve tubes not associated with companion cells have relatively thick walls and commonly are in direct contact with the vessels. The thick-walled sieve tubes have abundant cytoplasmic connections with contiguous vascular parenchyma cells; in contrast, connections between vascular parenchyma cells and thin-walled sieve tubes are rare. Connections are abundant, however, between the thin-walled sieve tubes and their companion cells; the latter have few connections with the vascular parenchyma cells. Plasmolytic studies on leaves of plants taken directly from lighted growth chambers gave osmotic potential values of about-18 bars for the companion cells and thin-walled sieve tubes (the companion cell-sieve tube complexes) and about-11 bars for the vascular parenchyma cells. Judging from the distribution of connections between various cell types of the vascular bundles and from the osmotic potential values of those cell types, it appears that sugar is actively accumulated from the apoplast by the companion cell-sieve tube complex, probably across the plasmalemma of the companion cell. The thick-walled sieve tubes, with their close spatial association with the vessels and possession of plasmalemma tubules, may play a role in retrieval of solutes entering the leaf apoplast in the transpiration stream. The transverse veins have chlorenchymatous bundle sheaths and commonly contain a single vessel and sieve tube. Parenchymatic elements may or may not be present. Like the thick-walled sieve tubes of the longitudinal bundles, the sieve tubes of the transverse veins have plasmalemma tubules, indicating that they too may play a role in retrieval of solutes entering the leaf apoplast in the transpiration stream.     

西瓜小叶脉超微结构研究

用透射电子显微技术研究了西瓜叶片小叶脉,结果表明,小叶脉是由大型维管束鞘细胞包围的维管束,维管束呈现大的头部和线形的柄部,柄部是单列细胞的木质部,由维管薄壁细胞和导管分子组成;头部是韧皮部,由维管薄壁细胞、伴胞和筛管分子组成。同一小叶脉内常见有超微结构特征显著不同的两种伴胞:一种伴胞体积小,与维管束鞘细胞接触面较小或不接触,细胞内有大液泡,细胞壁上没有胞间连丝或只有少数不分枝的胞间连丝,这种伴胞为2a型;另一种伴胞体积大,通常位于韧皮部两翼,不含大液泡而含大量小泡,与维管束鞘细胞接触面较大,接触面上有大量具分枝的胞间连丝,分枝部分比未分枝部分直径小,这种伴胞为中间细胞类型。显然,西瓜是小叶脉内兼具两种类型伴胞的植物。     

西瓜小叶脉超微结构研究

用透射电子显微技术研究了西瓜叶片小叶脉,结果表明,小叶脉是由大型维管束鞘细胞包围的维管束,维管束呈现大的头部和线形的柄部,柄部是单列细胞的木质部,由维管薄壁细胞和导管分子组成;头部是韧皮部,由维管薄壁细胞、伴胞和筛管分子组成。同一小叶脉内常见有超微结构特征显著不同的两种伴胞：一种伴胞体积小,与维管束鞘细胞接触面较小或不接触,细胞内有大液泡,细胞壁上没有胞间连丝或只有少数不分枝的胞间连丝,这种伴胞为2a型;另一种伴胞体积大,通常位于韧皮部两翼,不含大液泡而含大量小泡,与维管束鞘细胞接触面较大,接触面上有大量具分枝的胞间连丝,分枝部分比未分枝部分直径小,这种伴胞为中间细胞类型。显然,西瓜是小叶脉内兼具两种类型伴胞的植物。     

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.     

ULTRASTRUCTURE OF THE MATURE UNGERMINATED RICE (ORYZA SATIVA) CARYOPSIS. THE CARYOPSIS COAT AND THE ALEURONE CELLS

The results of a light and electron microscopic study of the caryopsis coat and aleurone cells in ungerminated, unimbibed rice (Oryza sativa) caryopses are presented. Surrounding the rice grain is the caryopsis coat composed of the pericarp, seed coat and nucellar layers. The outermost layer, the pericarp, consists of crushed cells and is about 10 μm thick. The seed coat, interior to the pericarp, is one cell thick and has a thick cuticle. Between the seed coat cuticle and endosperm are the remains of the nucellus. The nucellus is about 2.5 μm thick and has a thick cuticle adjacent to the seed coat cuticle. Interior to the caryopsis coat is the aleurone layer of the endosperm. The aleurone completely surrounds the rice grain and is composed of two cell types—aleurone cells that surround the starchy endosperm and modified aleurone cells that surround the germ. The aleurone cells of the starchy endosperm contain many aleurone grains and lipid bodies around a centrally located nucleus. The modified aleurone cells lack aleurone grains, have fewer lipid bodies than the other aleurone cells, and contain filament bundles (fibrils). Plastids of aleurone cells exhibit a unique morphology in which the outer membranes invaginate to form tubules and vesicles within the plastid. Transfer aleurone cells are not observed in the mature rice caryopsis.     

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     

Ultrastructure of minor veins inCucurbita pepo leaves

Summary The minor veins ofCucurbita pepo leaves were examined as part of a continuing study of leaf development and phloem transport in this species. The minor veins are bicollateral along their entire length. Mature sieve elements are enucleate and lack ribosomes. There is no tonoplast. The sieve elements, which are joined to each other by sieve plates, contain mitochondria, plastids and endoplasmic reticulum as well as fibrillar and tubular (190–195 diameter) P-protein. Fibrillar P-protein is dispersed in mature abaxial sieve elements but remains aggregated as discrete bodies in mature adaxial sieve elements. In both abaxial and adaxial mature sieve elements tubular P-protein remains undispersed. Sieve pores in abaxial sieve elements are narrow, lined with callose and are filled with P-protein. In adaxial sieve elements they are wide, contain little callose and are unobstructed. The intermediary cells (companion cells) of the abaxial phloem are large and dwarf the diminutive sieve elements. Intermediary cells are densely filled with ribosomes and contain numerous small vacuoles and many mitochondria which lie close to the plasmalemma. An unusually large number of plasmodesmata traverse the common wall between intermediary cells and bundle sheath cells suggesting that the pathway for the transport of photosynthate from the mesophyll to the sieve elements is at least partially symplastic. Adaxial companion cells are of approximately the same diameter as the adaxial sieve elements. They are densely packed with ribosomes and have a large central vacuole. They are not conspicuously connected by plasmodesmata to the bundle sheath.