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1.
Stem tissue of Lycopodium lucidulum Michx. was fixed in glutaraldehyde and postfixed in osmium tetroxide for electron microscopy. Although their protoplasts contain similar components, immature sieve elements can be distinguished from parenchymatous elements of the phloem at an early stage by their thick walls and correspondingly high population of dictyosomes and dictyosome vesicles. Late in maturation the sieve-element walls undergo a reduction in thickness, apparently due to an “erosion” or hydrolysis of wall material. At maturity, the plasmalemma-lined sieve elements contain plastids with a system of much convoluted inner membranes, mitochondria, and remnants of nuclei. Although the endoplasmic reticulum (ER) in most mature sieve elements was vesiculate, in the better preserved ones the ER formed a tubular network closely appressed to the plasmalemma. The sieve elements lack refractive spherules and P-protein. The protoplasts of contiguous sieve elements are connected with one another by pores of variable diameter, aggregated in sieve areas. As there is no consistent difference between pore size in end and lateral walls these elements are considered as sieve cells.  相似文献   

2.
Roots of Equisetum hyemale L. var. affine (Engelm.) A. A. Eat. were fixed in glutaraldehyde, postfixed in osmium tetroxide, and sieve elements of various ages were examined with the electron microscope. Young sieve elements are distinguished by their position within the vascular cylinder and by the presence of numerous refractive spherules, which originate within dilated portions of the endoplasmic reticulum (ER). Early in development, the sieve-element walls undergo a substantial increase in thickness. This is followed by the appearance of massive ER aggregates in the cytoplasm and then by a phase involving stacking and sequestering of the remaining ER. Nuclear degeneration is initiated shortly after the appearance of the ER aggregates. The chromatin condenses into masses of variable size along the inner surface of the nuclear envelope. The envelope then ruptures and chromatin is released into the cytoplasm. During the period of nuclear degeneration, mitochondria and plastids undergo structural modification, while components such as dictyosomes, microtubules, and ribosomes degenerate and disappear. The remaining cytoplasmic components assume a parietal position in the cell, leaving the lumen of the cell clear in appearance. At maturity, the plasmalemma-lined sieve element contains plastids, mitochondria, some ER, and refractive spherules. At this time many of the refractive spherules are discharged into the region of the wall. Pores between sieve elements occur largely on the end walls. During pore development, tubules of ER apparently traverse the pores, but because of the presence of massive callose deposits in the material examined, the true condition of mature pores could not be determined. The connections between mature sieve elements and pericycle cells are characterized by the presence of massive wall thickenings on the pericycle-cell side. Plasmodesmata in the wall thickening are matched by pores on the sieve-element side. Ontogenetic and cytoplasmic factors argue against use of the term “companion cell” for the vascular parenchyma cells associated with the sieve elements.  相似文献   

3.
The aerial shoots of Equisetum hyemale L. var. affine (Engelm.)A. A. Eat. were examined with the electron microscope as partof a continuing study of sieveelement development in the lowervascular plants. Young E. hyemale sieve elements are distinguishablefrom all other cell types within the vascular system by thepresence of refractive spherules, proteinaceous bodies whichdevelop within dilated portions of the endoplasmic reticulum(ER). Details of cell wall thickening differ between protophloemand metaphloem sieve elements. Following cell wall thickeningthe ER increases in quantity and aggregates into stacks. Shortlythereafter, nuclear degeneration is initiated. During the periodof nuclear degeneration some cytoplasmic components-dictyosomes,microtubules and ribosomes-degenerate and disappear, while organellessuch as mitochondria and plastids persist. The latter undergostructural modifications and become parietal in distribution.Eventually the massive quantities of ER are reduced, leavingthe lumen of the cell clear in appearance. At maturity the plasmalemma-linedsieve element contains a parietal network of tubular ER, aswell as mitochondria, plastids, and refractive sphemh At thistime many of the spherules are discharged into the region ofthe wall. Sieveelement pores occur in both lateral and end walls.At maturity many pores are traversed by large numbers of ERmembranes. The metaphloem sieve elements of the mid-internodalregions apparently are sieve-tube members. The connections betweenmature protophloem sieve elements and pericycle cells are associatedwith massive wall thickenings on the pericyclecell side.  相似文献   

4.
Leaf tissue of Isoetes muricata Dur. was fixed in glutaraldehyde and postfixed in osmium tetroxide for electron microscopy. The very young sieve elements can be distinguished from contiguous parenchyma cells by their distinctive plastids and the presence of crystalline and fibrillar proteinaceous material in dilated cisternae of the rough ER. During differentiation, the portions of ER enclosing this proteinaceous substance become smooth surfaced and migrate to the cell wall. Along the way they apparently form multivesicular bodies which then fuse with the plasmalemma, discharging their contents to the outside. At maturity, the sieve element contains an elongate nucleus, which consists of dense chromatin material, and remnants of the nuclear envelope. In addition, the mature sieve element is lined by a plasmalemma and a parietal, anastomosing network of smooth ER. Both plastids and mitochondria are present. P-protein is lacking at all stages of development. Tonoplasts are. not discernible in mature sieve elements. The end walls of mature sieve elements contain either plasmodesmata or sieve pores or both, but only plasmodesmata occur in the lateral walls.  相似文献   

5.
Root tissues of Isoetes muricata Dur. were fixed in glutaraldehydeand postfixed in osmium tetroxide for electron microscopy. Veryyoung root sieve elements can be distinguished from contiguousparenchyma cells by the presence of crystalline and/or fibrillarproteinaceous material in dilated cisternae of rough endoplasmicreticulum (ER). Similar crystalline-fibrillar material accumulatesin the perinuclear space. During differentiation, the portionsof ER enclosing this proteinaceous substance become smooth surfacedand migrate to the cell wall. Along the way many of them formmultivesicular bodies which fuse with the plasmalemma, dischargingtheir contents toward the wall. Nuclear degeneration is pycnotic.At maturity, the sieve element contains a degenerate, filiformnucleus, plastids, and mitochondria. In addition, the wall ofthe mature sieve element is lined by a plasmalemma and a parietalnetwork of smooth ER. Sieve-area pores are present in both endand lateral walls of mature sieve elements. Whereas a singlecluster of pores occurs in each end wall, the pores of the lateralwalls are solitary and few in number.  相似文献   

6.
Corm tissue of Isoetes muricata Dur. was fixed in glutaraldehyde and postfixed in osmium tetroxide for electron microscopy. Very young secondary sieve elements can be distinguished from contiguous cambial cells by their distinctive plastids and by the presence of crystalline and/or fibrillar proteinaceous material in dilated cisternae of rough endoplasmic reticulum (ER). At maturity, the sieve elements are lined by the plasmalemma and a parietal, anastomosing network of smooth ER. Degenerate nuclei persist in all mature sieve elements. In addition, mature sieve elments contain plastids and mitochondria. Sieve-area pores are present in all walls. The lateral meristem of I. muricata consists of 2–3 layers of cells year-round. Judging from numerous collections made between October 1972 and July 1975, new sieve-element differentiation precedes cambial activity by about a month. Early in May, 1–2 cells immediately adjacent to already mature sieve elements differentiate directly into sieve elements without prior division. In early June, at about the time sieve-element differentiation is completed, cambial division begins. Division is sporadic, not uniform throughout the meristem. Dormancy callose accumulates in the secondary sieve elements in late October, and is removed in early May, at about the same time new sieve-element differentiation begins. Cells of the dormant cambium are characterized by the presence of numerous small vacuoles and large quantities of storage materials, including lipid droplets, starch grains, and tannin. By contrast, active cambial cells contain few large vacuoles with little or no tannin, and they have little storage material.  相似文献   

7.
The differentiation of sieve elements from inner cortical cells of the stipe of Laminaria saccharina (L.) Lamour. involves the development of a well-structured protoplast and an end wall possessing evenly spaced pores which are visualized by electron microscopy. The protoplast consists of organelles which are commonly found in brown algal cells, including nuclei, cup- or horseshoe-shaped chloroplasts, dictyosomes, mitochondria, and ER. Mitochondria and clusters of small vacuoles, presumably redistributed by the surging effect which occurs in sieve elements, were routinely observed in the vicinity of the end wall. Chloroplasts were seen in progressively degenerated states in older sieve elements, yet nuclei were determined to be non-necrotic. Numerous pores along the end walls interconnect adjacent sieve elements. Each pore is traversed by a strand of cytoplasm and surrounded by plasmalemma. The pores are open and possess no callose. In this paper the sieve element ultrastructures of L. saccharina are compared to those in L. groenlandica, Alaria marginata, Nereocystis lutkeana and Macrocystis pyrifera, and a possible phylogenetic specialization of sieve elements is presented in table form and discussed.  相似文献   

8.
The structure of the mature leaves of 13 species from 9 families of homosporous leptosporangiate ferns was examined by light and electron microscopy. In 11 species (Adiantum pedatum L., Athyrium angustum Roth., Cyathea dregei Sm., Lygodium palmatum Sw., Mohria caffrorum (L.) Desv., Oleandra distenta Kuntae, Pellaea calomelanos (Sw.) Link, Pityrogramma calomelanos (L.) Link var. austro-americana (Domn.) Farw., Trichomanes melanotrichum Schlechtend., Vittaria guineensis Desv., and Woodwardia orientalis Sw.) the lamina veins are collateral; in two (Phlebodium aureum and Platycerium bifurcatum), bicollateral as well as collateral veins are present. The vascular bundles in the midribs of C. dregei and those in the petioles and midribs of Phlebodium and Platycerium are concentric. All of the vascular bundles in the homosporous leptosporangiate ferns studied are delimited by a tightly arranged cylinder of endodermal cells with Casparian strips. Within the veins without parenchymatic xylem sheaths, some sieve elements commonly abut tracheary elements with hydrolyzed primary walls. The majority of vascular parenchyma cells contact both sieve elements and tracheary elements, although some parenchyma cells are associated with only one type of conducting cell. Transfer cells (parenchyma cells with wall ingrowths) occur in the veins of 6 species examined. Most of the vascular parenchyma cells, however, have no distinctive structural characteristics. The sieve elements of the homosporous leptosporangiate ferns are very similar structurally and each consists of a plasmalemma, a parietal, anastomosing network of smooth endoplasmic reticulum (ER), and variable numbers of refractive spherules, plastids and mitochondria. The sieve elements of L. palmatum also contain plasmalemma tubules. The parenchymatic cells of the leaf (mesophyll, endodermal and vascular parenchyma cells) are united by desmotubule-containing plasmodesmata. The sieve elements are connected to each other by sieve pores and to parenchymatic cells by pore-plasmodesma connections. The sieve-area pores contain variable amounts of membranous material, apparently ER membranes, but do not occlude them. These membranes commonly are found in continuity with the parietal ER of the lumen. Based upon the relative frequencies of cytoplasmic connections between cell types, the photosynthates may move from the mesophyll to the site of phloem loading via somewhat different pathways in different species of homosporous leptosporangiate ferns.  相似文献   

9.
The structure and differentiation of the sieve element of lower vascular plants is reviewed using data obtained primarily from ultrastructural investigations conducted during the last ten years. During the last decade the phloem of representatives from every major group of the ferns and fern allies has been examined with the electron microscope and from these studies a rather clear picture has emerged of the structure of the sieve element protoplast in this diverse group of plants. Present data indicate that although the details of sieve-element differentiation may differ, the protoplasts of the mature sieve elements in the various groups of lower vascular plants are remarkably similar in structure. Each consists of a plasmalemma, a parietal, anastomosing network of smooth ER, plastids, mitochondria and, with the exception of the lycopods, variable numbers of refractive spherules. The protoplasts of mature sieve elements are joined by plasmalemma-lined connections, each arising from a single plasmodesma during the course of sieve element differentiation. The size of the connections in the mature elements range from plasmodesmata-like structures to relatively wide sieve-area pores, depending on the species. Moreover, the contents of the cytoplasmic connections vary somewhat according to the species. Whereas in the lycopods, the sieve-area pores are virtually unoccluded by any cytoplasmic material, the cytoplasmic connections of all other lower vascular plants examined with the electron microscope contain variable amounts of membranous material, apparently tubular elements of ER. In Equi-setum hyemale, Psilotum nudum and the eusporangiate and protoleptosporangiate ferns, the ER membranes are very numerous and virtually occlude the pores. Furthermore, the membranes apparently are not connected with the parietal ER in the lumen of the cell. The sieve-area pores of the leptosporangiate ferns also contain ER membranes, however, they are not as abundant as the membranes of the eusporangiate and protoleptosporangiate ferns. In addition, in the leptosporangiate ferns the pore membranes apparently are united with the parietal ER in the lumen of the cell.  相似文献   

10.
The sieve elements of Nereocystis from the base of phylloids contain numerous small vesicles, cytoplasm, ribosomes, and the usual organelles and membrane systems, including nuclei, plastids, mitochondria, dictyosomes, and endoplasmic reticulum. They have a thick secondary wall layer which is deposited along the longitudinal walls and at the sieve plate excluding the sieve pores. The sieve pores range in diameter from 100 to 400 nm and are lined by plasmalemma. The sieve elements from the hollow basal parts of the pneumatocyst show essentially the same features but have larger and fewer vesicles, relatively little cytoplasm, larger sieve pores, 400–900 nm in diameter, and may lack a nucleus. In old sieve elements there are large deposits of callose on the sieve plate and along the longitudinal wall; the vesicles seem to break down, and the protoplast appears necrotic. It is concluded that the trumpet hyphae and sieve tubes are basically the same type of cell, and that the trumpet-shape of the sieve elements is due to their passive stretching during extension growth of the organ in which they occur. There are minor but significant differences among the sieve elements from different regions of the thallus which may reflect possible levels of structural specialization of the sieve elements within the same plant.  相似文献   

11.
Sieve cell differentiation in the primary phloem of Ephedra viridis is first indicated by an increase in thickness of the wall, which begins in the corners of the cell, and next by the proliferation of smooth tubular endoplasmic reticulum (ER). As differentiation proceeds, cisternae of rough ER form stacks along the wall, losing their ribosomes in the process. Concomitantly, all of the mitochondria, plastids, and ER become parietal in distribution, and the vacuoles collapse. Nuclear degeneration is pycnotic and accompanied by the formation of tubular invaginations of the nuclear envelope into the peripheral chromatin. At maturity, an anastomosing network of smooth ER borders the plasmalemma, interconnecting aggregates of smooth tubular ER located primarily opposite the sieve areas. In addition to ER, the mature sieve cell contains mitochondria, plastids, and remnants of the degenerate nucleus, all of which are parietal in distribution. P-protein is lacking at all stages of sieve cell development. Sieve pore and compound median cavity development is similar to that reported for the sieve cells of conifers. Albuminous cells are associated with the sieve cells of the metaphloem throughout the shoot but with sieve cells of the protophloem only in the node. Among their cytoplasmic components are broad bundles of microfilaments spatially associated with a complex system of rough and smooth ER.  相似文献   

12.
As part of a continuing study of sieve elements in primitive angiosperms, a study of this cell type was undertaken in Liriodendron tulipifera. A typical ontogenetic sequence was observed in which synthetic processes such as wall thickening are followed in time by cellular lysis of nucleus, ribosomes, microtubules, vacuoles, and dictyosomes. This lysis is selective in that certain cellular components (e.g., the plasmalemma) remain unaffected. Concomitant with lysis is the formation of sieve-area pores from plasmodesmata. Comparison of pore size on end and lateral walls indicates that the use of the term “sieve tube” rather than “sieve cell” to describe these elements is appropriate.  相似文献   

13.
Both intact and cut duckweed plants were prepared for electron microscopy. Plants which are prepared intact do not exhibit callose formation during development of sieve-plate pores. Future pore sites can be recognized by the presence of median cavities that are unassociated with callose platelets. These cavities are first seen in the region of the compound middle lamella and are lined by a plasmalemma. As end walls thicken, the cavities increase in size until open pores of uniform width are formed. Mature sieve plates of intact-prepared plants are also devoid of callose. Fully opened pores are lined by a plasmalemma and are only traversed by an occasional tubule of endoplasmic reticulum. Plants which have been cut prior to fixation possess mature sieve plates containing callose. The pores of developing sieve plates in cut plants exhibit small amounts of callose. Except for the lack of callose, lateral wall connections between sieve elements and contiguous cells are similar in development and mature state to those reported for other species.  相似文献   

14.
This report presents a combined investigation of ultrastructural and enzymatic changes in the procambium from late winter to early spring. In January the procambial cells of dormant Salix buds have a convoluted plasma membrane with many plasmalemmasomes, numerous lipid bodies, large stacks of rough ER and plastids surrounded by smooth ER profiles. Several small lysosomes show activity of ATPase and acid phosphatases. In addition ER, nuclear envelopes, dictyosomes, and thylakoids have ATPase activity, and ER and plasmalemma, and nuclei also show acid phosphatase activity. In February metabolism seems to increase as indicated by lysosomes with membranous formations, dilated ER, nuclear envelopes, spiny vesicles, and polysomes. ATPase activity occurs in plasmalemma and vacuoles, and acid phosphatases in the middle lamella region of walls, in plasmalemma, vacuoles, ER, and nuclei. At the end of March, when growth starts inside the buds, but before they break, the stacks of rough ER disappear, and the vacuoles coalesce. Most of the lipid bodies have disappeared and the plastids have accumulated starch. Cell division and differentiation of procambial cells to protophloem and protoxylem have started. The distribution of ATPase increases; activity is found in walls and plasmalemma, and only a few small vacuoles still have ATPase and acid phosphatase activity. Notable is the appearance of ATPase in mitochondrial cristae and nucleoli and the occurrence of rather high levels also in endomembranes and dictyosomes.  相似文献   

15.
Sieve elements of various ages were examined in petioles and midribs of Platycerium bifurcatum (Cav.) C. Chr. and Phlebodium aureum (L.) J. Sm., only older ones in similar parts of leaves of Polypodium schraderi Mett. and Microgramma lycopodioides (L.) Copel. Nacreous walls apparently are formed by most, if not all, protophloem and metaphloem sieve elements in all four species. In Platycerium and Phlebodium nacreous wall formation is closely correlated with the appearance of numerous membranes or vesicles in the region of the wall. These extracytoplasmic membranes apparently are derived from protrusions of the plasmalemma. After the nacreous layer is fully thickened, many endoplasmic reticulum (ER) membranes apparently end up outside the plasmalemma of Platycerium, where they degenerate and gradually intergrade in appearance with the fibrillar material comprising the nacreous thickening. In Phlebodium, Polypodium, and Microgramma the ER forms multivesicular bodies. As the cells approach maturity, the membranes delimiting the multivesicular bodies fuse with the plasmalemma and their vesicular contents, which are not discharged into the region of the wall, disappear. Gradually, the nacreous layer decreases in thickness and disappears. At maturity the enucleate sieve-element protoplasts of all four species are essentially similar. They are lined by a plasmalemma and a parietal, anastomosing network of ER and contain both plastids and mitochondria. The plastids in Polypodium and Microgramma are chloroplasts, but those in Platycerium and Phlebodium lack grana and intergrana lamellae.  相似文献   

16.
The physiological phloem equivalents, leptoids, of the polytrichaceous moss Atrichum undulatum appear to be similar to the nacreous sieve elements that occur in many higher plants. These leptoids are elongated cells with nacreous thickenings on their radial and tangential walls. Their oblique end walls, which lack such thickenings, are traversed by numerous pores through which the plasmalemma, endoplasmic reticulum, and cytoplasm are continuous between adjacent leptoids of a longitudinal file. These end walls closely resemble the simple sieve areas of the sieve elements found in Polypodium vulgare. The leptoid sieve pores have a median expanded area and frequently are occluded by small amorphous protein plugs at each end. Also, callose was observed as electron-luscent areas both on the faces of the end walls and as a thin cylinder surrounding the lateral area of each pore. Amorphous and granular cytoplasmic contents of the leptoids appear to be morphologically similar to the slime (P-protein) found in the sieve-tube elements of many angiosperms. Differentiating leptoids are characterized by the formation of numerous membrane-bound protein bodies in close association with polysomes and endoplasmic reticulum. As the leptoid matures, the contents of the protein bodies become dispersed in the cytoplasm. Ultrastructurally and ontogenetically the leptoids in the gametophores of A. undulatum appear almost identical to the sieve elements of P. vulgare and therefore should be considered sieve elements rather than phloem-like equivalents.  相似文献   

17.
Summer and winter (July and January) samples of secondary phloem of Tilia americana were studied with the electron microscope. Parenchyma cells contain: nuclei, endoplasmic reticulum, ribosomes, plastids, mitochondria and occasional dictyosomes. Well-defined tonoplasts separate vacuoles from cytoplasmic ground substance. Vacuoles often contain tannins. Lipid droplets are common in cytoplasm. Endoplasmic reticulum–connected plasmodesmata are aggregated in primary pit fields. Companion cells differ from parenchyma cells in having numerous sieve-element connections, possibly slime, and in lacking plastids. Mature, enucleate sieve elements possess 1–4 extruded nucleoli. Numerous vesicles occupy a mostly parietal position in association with plasmalemma. The mature sieve element lacks endoplasmic reticulum, organelles (except for few mitochondria) and tonoplast. In OsO4– and glutaraldehyde-fixed elements, slime has a fine, fibrillar appearance. Normally, these fine fibrils are organized into coarser ones which form strands that traverse the cell and the plasmalemma-lined pores of sieve plates and lateral sieve areas.  相似文献   

18.
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.  相似文献   

19.
Summary Phloem tissues ofMicrosorium scolopendria (Polypodiaceae) andPsilotum nudum (Psilotaceae) were examined with light and electron microscopes. The characteristic refractive spherules in the sieve elements ofM. scolopendria apparently develop from endoplasmic reticulum-derived cytoplasmic vesicles. In both taxa they have not been observed to be spatially related to plastids or mitochondria. Refractive spherules contain protein and often occur in the peripheral cytoplasm of mature sieve elements. InM. scolopendria they also occur in pericycle cells. Significant differences in refractive spherule substructure occur between the two taxa studied.  相似文献   

20.
Both thick- and thin-walled sieve tubes in leaf-blade veins of Hordeum vulgare L. exhibit a distinct, electron-opaque inner wall layer after fixation in glutaraldehyde-osmium tetroxide and staining with uranyl acetate and lead citrate. This inner wall layer is thickest at the sieve plates and lateral sieve areas where it is permeated by a labyrinth of tubules formed by the plasmalemma. Along the lateral walls between sieve areas the inner wall layer apparently is penetrated by numerous microvilli-like evaginations of the plasmalemma, giving the cell wall-plasmalemma interface the appearance of a brush border. It is suggested that a similar brush-border-like structure may occur at the cell wall-plasmalemma interface of sieve elements in a wide variety of vascular plants.Abbreviation ER endoplasmic reticulum  相似文献   

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