STUDIES ON THE LEAF OF POPULUS DELTOIDES (SALICACEAE): QUANTITATIVE ASPECTS,AND SOLUTE CONCENTRATIONS OF THE SIEVE-TUBE MEMBERS

The vascular system of the leaf of Populus deltoides Bartr. ex Marsh, was examined quantitatively, and plasmolytic studies were carried out to determine the solute concentrations of sieve-tube members at various locations in the leaf. Both the total number and total crosssectional area of each cell type decreases with decreasing vein size. Although the proportion of phloem occupied by sieve tubes varies considerably from location to location, a linear relationship exists between cross-sectional area of the vascular bundles and both total and mean cross-sectional area of sieve tubes. Collectively, the cross-sectional area of all tertiary and minor veins feeding into a secondary exceeds the total cross-sectional area of sieve tubes at the base of that secondary. Moreover, the total volume of sieve tubes in the “catchment area” of a secondary vein is much greater than the total sieve tube volume of the secondary itself. Both tracheary elements and sieve-tube members undergo a reduction in both total and mean crosssectional area in the constricted zone at the base of the leaf. The plasmolytic studies revealed the presence of positive concentration gradients in sieve tubes of the lamina from the minor veins and tips of the secondaries to the bases of the secondaries and their associated subjacent midvein bundles and from the upper to lower portions of the median bundle of the midvein.     

STUDIES ON THE LEAF OF AMARANTHUS RETROFLEXUS (AMARANTHACEAE***): MORPHOLOGY AND ANATOMY

The leaf of Amaranthus retroflexus L. was examined with the light microscope to determine its vasculature and the spatial relationship of the vascular bundles to the mesophyll. Seven leaf traces enter the petiole at the node and form an arc that continues acropetally in the petiole as an anastomosing system of vascular bundles. Upon entering the lamina, the arc of bundles gradually closes and forms a ring of anastomosing bundles that constitutes the primary vein, or midvein, of the leaf. As the midvein progresses acropetally, branches of the bundles nearest the lamina diverge outward and continue as secondary veins toward the margin on either side of the lamina. Along its course the midvein undergoes a gradual reduction in number of bundles until only one remains as it approaches the leaf tip. Tertiary veins arise from the secondaries, and minor veins commonly arise from all orders of major veins, as well as from other minor veins. All of the major veins are associated with rib tissue, although the ends of the tertiaries may resemble minor veins, which are completely encircled by chlorenchymatic bundle sheaths and mesophyll cells that radiate out from the sheaths. A specialized minor vein, the fimbrial vein, occurs just inside the margin of the leaf. Most of the mesophyll cells—the so-called “Kranz mesophyll cells”—are in direct contact with the bundle sheaths, but some—the so-called “nonKranz mesophyll cells”—lack such contact. Non-Kranz mesophyll cells are especially prominent where they form a network of mostly horizontally oriented cells just above the lower epidermis. Guard cells of both the upper and lower epidermis are spatially associated with nonKranz mesophyll cells.     

VASCULAR SYSTEM OF LODICULES OF DACTYLIS GLOMERATA L.

The vascular system for the two lodicules in a floret of Dactylis glomerata L. was studied in serial sections. The floret stele contained a few modified tracheary elements and xylem transfer cells enveloped by a phloem of squat sieve-tube members and intermediary cells. A single sieve tube and associated phloem parenchyma exited the right and left sides of the stele and upon nearing the base of each lodicule branched and formed the minor veins of the lodicule. The minor veins underwent limited branching and anastomosing to form a small three-dimensional system which described an arc during its ascent in the adaxial portion of each lodicule. The sieve tubes in the minor veins extended halfway up the lodicule and contained short sieve-tube members with transverse, slightly oblique, or lateral simple sieve plates. The associated phloem parenchyma cells were intermediary cells, companion cells, and less intimate parenchyma cells. Intermediary cells terminated the minor veins and touched the distal ends of the terminal sieve-tube members, which lacked distal sieve plates. Although the transverse area of the sieve-tube members remained constant up the lodicule, the transverse area of the associated phloem parenchyma fluctuated.     

STUDIES ON THE LEAF OF POPULUS DELTOIDES (SALICACEAE): MORPHOLOGY AND ANATOMY

Mature field- and growth-chamber-grown leaves of Populus deltoides Bartr. ex Marsh. were examined with light and scanning electron microscopes to determine their vasculature and the spatial relationships of the various orders of vascular bundles to the mesophyll. Three leaf traces, one median and two lateral, enter the petiole at the node. Progressing acropetally in the petiole these bundles are rearranged and gradually form as many as 13 tiers of vascular tissue in the petiole at the base of the lamina. (Most leaves contained seven vertically stacked tiers.) During their course through the midrib the tiers “unstack” and portions diverge outward and continue as secondary veins toward the margin on either side of the lamina. As the midvein approaches the leaf tip it is represented by a single vascular bundle which is a continuation of the original median bundle. Tertiary veins arise from the secondary veins or the midvein, and minor veins commonly arise from all orders of veins. All major veins–primaries, secondaries, intersecondaries, and tertiaries–are associated with rib tissue, while minor veins are completely surrounded by a parenchymatous bundle sheath. The bundle sheaths of tertiary, quaternary, and portions of quinternary veins are associated with bundle-sheath extensions. Minor veins are closely associated spatially with both ad- and abaxial palisade parenchyma of the isolateral leaf and also with one or two layers of paraveinal mesophyll that extend horizontally between the veins. The leaves of growth-chamber-grown plants had thinner blades, a higher proportion of air space, and greater interveinal distances than those of field-grown plants.     

STUDIES ON THE LEAF OF POPULUS DELTOIDES (SALICACEAE): ULTRASTRUCTURE,PLASMODESMATAL FREQUENCY,AND SOLUTE CONCENTRATIONS

The minor veins and contiguous tissues of mature leaves of Populus deltoides Bartr. ex Marsh. were examined with the electron microscope to determine the ultrastructural characteristics of the component cells and to determine the structure, distribution, and frequency of plasmodesmata between the various cell types. In addition, plasmolytic studies were carried out to determine the solute concentrations of the various cell types of the minor veins and contiguous tissues. The cells comprising the mesophyll and bundle sheath contain all the components typical of photosynthetic cells. Paraveinal mesophyll cells and bundle-sheath cells have fewer microbodies and smaller chloroplasts than do palisade parenchyma cells. Vascular parenchyma and companion cells tend to intergrade with one another structurally but can be distinguished from one another by their characteristic plastids. The mature, enucleate sieve-tube member is lined by a parietal layer of cytoplasm consisting of plasmalemma, endoplasmic reticulum, mitochondria, plastids, and P-protein. Plasmodesmata occur along all possible routes from the palisade parenchyma cells to the sieve tubes of the minor veins, and their frequency increases with increasing proximity to the sieve-tube members. Plasmolytic studies revealed that the paraveinal mesophyll cells had a higher C₅₀ (estimated mannitol concentration plasmolyzing, on the average, 50% of a given cell type) than any other cell type of the leaf. Concentration gradients existed along the palisade cell/bundle-sheath cell/companion cell (or vascular parenchyma cell) route as well as along the paraveinal mesophyll cell/bundle-sheath cell/companion cell (or vascular parenchyma cell) route. Considering the frequency of plasmodesmata along these routes, it is conceivable that photosynthate diffuses from palisade cells to the companion cells along concentration gradients. Within the minor veins, the C₅₀ was higher for sieve-tube members than for either companion cells or vascular parenchyma cells, indicating that loading of the sieve tubes is an active, energy-dependent process.     

MORPHOLOGY AND ANATOMY OF THE LEAF OF COLEUS BLUMEI (LAMIACEAE)

Leaves of the Princeton and a variegated clone of Coleus blumei Benth. were examined with the light microscope to determine the course of their vasculature and the spatial relationship between the mesophyll, bundle sheath, and vascular tissues. In Princeton clone leaves two leaf traces enter the petiole at the node and quickly branch to form an arc of bundles which undergo further divisions as well as fusions in the distal half of the petiole. The anastomosing arc of bundles reaches its greatest complexity in the base of the midvein, where its lateral-most bundles unite and diverge outward to form secondary veins. As the midvein bundles continue acropetally, they gradually fuse more and divide less until only a single bundle remains, from which secondaries and smaller veins branch. Major (ribbed) veins include not only the midvein and secondaries but also tertiary and quaternary veins. Decreasing vein size is accompanied by increasing direct contact between vascular and photosynthetic tissues. Minor veins, which make up 86% of the total vein length, are completely surrounded by photosynthetic bundle sheaths and mesophyll consisting of palisade and spongy parenchyma. Statoliths occur in a layer of cells just outside the phloem of the petiole-midrib axis and secondary veins. Functional hydathodes are present at the apices of the marginal teeth. The overall organization of tissues in variegated leaves differs little in either the green or albuminous areas from corresponding (but always green) regions of Princeton leaves. Chloroplasts are lacking in mesophyll, bundle-sheath, and most guard cells of the albuminous region but are present in guard cells which are within 1 mm of green areas.     

Studies onArtemisia afra Jacq.: The phloem in stem and leaf

Summary The structure of the phloem was studied in stem and leaf ofArtemisia afra Jacq., with particular attention being given to the sieve element walls. Both primary and secondary sieve elements of stem and midvein have nacreous walls, which persist in mature cells. Histochemical tests indicated that the sieve element wall layers contained some pectin. Sieve element wall layers lack lignin. Sieve elements of the minor veins (secondary and tertiary veins) lack nacreous thickening, although their walls may be relatively thick. These walls and those of contiguous transfer cells are rich in pectic substances. Transfer cell wall ingrowths are more highly developed in tertiary than in secondary veins.     

HISTOGENESIS OF LEAF VENATION IN TRIFOLIUM WORMSKIOLDII (LEGUMINOSAE)

The histogenesis of the leaf venation in Trifolium wormskioldii was followed, using whole cleared leaflets and transverse and paradermal microtome sections. Mature leaflets are about 8 mm in length. The midvein precambium appears in leaflets 100 μ in length; secondary vein procambium is first seen in 230 μ leaflets; procambium of minor veins first appears in leaflets 400-600 μ in length. Phloem is apparent before xylem, in midvein, secondary veins, and minor veins. Vein endings are initiated and they mature last. No evidence was found to support the theory of “vein breakage.” In the greenhouse 24 days elapsed from leaf initiation to exposure of leaf tips; 6 more days elapsed until full leaf expansion.     

贵州小檗属植物(小檗科)叶脉序研究

采用制作叶脉标本和透明叶标本的方法,对贵州产28种2变种小檗属植物叶脉特征进行比较研究。结果表明:贵州小檗属植物的脉序类型有5种:半达缘羽状脉、花环状半达缘羽状脉、简单弓形羽状脉、花环状弓形羽状脉和混合型。叶脉分支一般有五级:1一级脉构架均为羽状脉,粗度有很粗、粗、中等粗细和纤细四种类型,分支方式包括单轴分支和合轴分支;2粗二级脉构架中有分支达缘或分支均不达缘,与中脉夹角变化各异,内二级脉存在或缺失,细二级脉半达缘、真曲行或简单弓形,间二级脉类型复杂多变但频度种间有差异;3三级脉贯串型、结网型或分支型;4四、五级脉网状或自由分支且常混合在一起。脉间区从发育差到良好,小脉从不分支到不均等分支等各种类型均有,叶缘末级脉缺失、不完整、钉状和环状。大部分种类叶缘具齿,每1cm齿数目和齿内腺点的特性等特征在不同种类间有区别,具有鉴定价值,但齿其它特征复杂多变或种间区别较小,同时齿内脉性状也不稳定。此外,齿的有无会对脉序类型产生影响。小檗属植物叶脉类型存在种间差异,具有重要的分类学价值,叶脉类型的变化和复杂程度显示了该属植物的进化特点;叶齿的有无和齿特征具有分类学和系统学意义。基于叶脉特征的研究结果并结合重要的外部形态学特征编制了贵州小檗属植物的分种检索表。研究结果可为小檗属植物分类寻找新的依据并探讨其系统学意义。     

Quantitative survey of sieve tube distribution in foliar terminal veins of ten dicot species

Quantitative data on sieve tubes in foliar terminal veins (vein endings) were added to the meager published information from only five dicot species. Correlations with other minor vein configurations were also explored. Leaf samples from ten species of dicots (Oxalis nelsonii, O. pes-capri, O. rubra, O. stricta [Oxalidaceae], Caesalpinia pulcherrima. Glycine max, Trifolium repens [Leguminosae], Ampelamus albidus [Asclepidaceae], Eupatorium rugosum [Asteraceae], and Polygonum convolvulus [Polygonaceae]) were selected for two quantitative procedures: 1) a survey of the arrangement of terminal veins and distribution of sieve tubes in terminal veins in 100 areoles per species using stained leaf clearings; and 2) a search for correlations of sieve tube distribution with number and branching patterns of terminal veins, and with sizes of areoles using image analysis. Two Oxalis species (O. pes-capri and O. stricta) had the smallest areoles and virtually no sieve tubes in any terminal vein. Polygonum convolvulus, at the other extreme, had sieve tubes extending to the tips of most terminal veins. The other species had various intermediate sieve tube configurations. The data indicate that species with few or no sieve tubes associated with their terminal veins, regardless of the number of terminal veins per areole, have smaller areoles. These results may have implications regarding the entry of leaf photosynthates into the vascular system.     

Leaf structure and translocation in sugar beet

Anatomical and ultrastructural details of a translocating 10-cm leaf of sugar beet (Beta vulgaris L. var. Klein Wanzleben) were correlated with translocation rate data. The minor veins were found to be 13 times as extensive as the major veins and measure 70 cm/cm² leaf lamina. Measurements disclosed that a 33-μ length of minor vein services 29 mesophyll cells with the result that translocate moves an average of 73 μ or 2.2 cell diameters during transport from mesophyll cells to a minor vein. High-resolution, freeze-dry autoradiography revealed that assimilates accumulate in organelle-rich cells of the minor vein phloem. Correlation of phloem volume and loading rate for minor veins yielded an uptake rate of 735 μmoles of sucrose per g fresh weight of phloem. The arrangement and structural features of minor veins appeared to be consistent with the concept that vein loading precedes translocation.     

A QUANTITATIVE STUDY OF SIEVE-TUBE DIFFERENTIATION IN VEGETATIVE SHOOT APICES OF COLEUS

Quantitative methods, with round-the-clock collecting of large samples in successive years, have uncovered several new phenomena of sieve-tube differentiation in young leaves of Coleus vegetative shoots. In small leaves (1–350 μ), there are no sieve tubes in the leaf itself, but they differentiate acropetally in the two traces to each leaf. Regression lines fitted to the data for leaf length vs. most distal position of sieve tubes in the traces support the view that differentiation is steady and acropetal, but they also reveal that differentiation in the traces falls steadily farther behind elongation of the leaf. Leaves more than 500 μ long have sieve tubes close to their tips. An intensive search of leaves of intermediate lengths revealed an isolated locus of sieve-tube differentiation. These relationships were reproducible year after year. Every plant with discontinuous strands of sieve tubes in the second leaf pair had discontinuous xylem in the third. This isolated locus was not seen before, probably because of small samples and daytime collections; most of our cases were from night collections.     

Evidence for two pathways of phloem loading

The minor veins of small leaf discs, punched out of mature leaves and incubated in ¹⁴C-sucrose solution, appear labeled in macro- and microautoradiographs. Discs with a labeled vein pattern and with labeled sieve tubes in microautoradiographs were found in Beta vulgaris, Vicia faba, Gomphrena globosa and Antirrhinum majus . However, in several other plant species, minor veins appeared unlabeled in macroautoradiographs when the discs were incubated in ¹⁴C-sucrose. Mesophyll cells ( Acer pseudoplatanus, Juglans regia, Fagia, sylvatica, Syringa vulgaris, Laburnum anagyroides ), bundle-sheath cells of major veins ( Salix viminalis, Robinia pseudoacacia, Commelina communis ) or epidermal layers ( Ginkgo biloba, Chlorophytum comosum ) appeared labeled. Lack of radioactivity in sieve tubes of this latter group was confirmed by microauto-radiography. Using ¹⁴C-glucose instead of ¹⁴C-sucrose, leaf discs of Beta vulgaris showed no labeled vein pattern and in microautoradiographs the sieve tubes appeared unlabeled. In view of the by-pass phloem loading, this study provides evidence for two pathways of phloem loading.     

MORPHOLOGY AND VASCULATURE OF THE LEAF OF POTATO (SOLANUM TUBEROSUM)

The leaf and stem of the potato plant (Solanum tuberosum L. cv. Russet Burbank) were studied by light microscopy to determine their morphology and vasculature; scanning electron microscopy provided supplemental information on the leaf's morphology. The morphology of the basal leaves of the potato shoot is quite variable, ranging from simple to pinnately compound. The upper leaves of the shoot are more uniform, being odd pinnate with three major pairs of lateral leaflets and a number of folioles. The primary vascular system of the stem is comprised of six bundles, three large and three small ones. The three large bundles form a highly interconnected system through a repeated series of branchings and arch-producing mergers. Two of the three large bundles give rise to short, lateral leaf traces at each node. Each of the small bundles in the stem is actually a median leaf trace which extends three internodes before diverging into a leaf. The three leaf traces enter the petiole through a single gap; thus the nodel anatomy is three-trace unilacunar. Upon entering the petiole, each of the laterals splits into an upper and a lower lateral. Whereas the upper laterals diverge entirely into the first pair of leaflets, the lower laterals feed all of the lateral leaflets through a series of bifurcations. Prior to their entering the terminal leaflet, the lower laterals converge on the median bundle to form a single vascular crescent which progresses acropetally into the terminal leaflet as the midvein, or primary vein. In the midrib, portions of the midvein diverge outward and continue as secondaries to the margin on either side of the lamina. Near the tip of the terminal leaflet, the midvein consists of a single vascular bundle which is a continuation of the median bundle. Six to seven orders of veins occur in the terminal leaflet.     

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.     

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.     

LEAF OF SPINACIA OLERACEA (SPINACH): ULTRASTRUCTURE,AND PLASMODESMATAL DISTRIBUTION AND FREQUENCY,IN RELATION TO SIEVE-TUBE LOADING

Minor veins and contiguous tissues of the Spinacia oleracea leaf were analyzed by electron microscopy to determine the characteristics of the component cells and the structure, distribution, and frequency of plasmodesmata between the various cell types of the leaf. Mesophyll and bundle-sheath cells contain components typical of photosynthetic cells although the latter cell type contains smaller chloroplasts and fewer mitochondria and microbodies than the mesophyll cells. In addition, the mesophyll cells contain numerous invaginations of the plasmalemma bordering the chloroplasts and evaginations of the outer membrane of the opposing chloroplast envelope. In places, these membranes appear continuous with each other. The minor veins consist of tracheary elements, xylem parenchyma cells, sieve-tube members, companion and phloem parenchyma cells, and other cells simply designated vascular parenchyma cells. The companion and phloem parenchyma cells are typically larger than the sieve-tube members with the companion cells containing a much denser cytoplasm that the phloem parenchyma. Cytoplasmic connections occur along all possible routes from the mesophyll to the sieve-tube members and consist of either simple or branched plasmodesmata between parenchymatic elements or pore-plasmodesmata between the sieve-tube members and parenchyma cells. The highest frequency of plasmodesmata occurs between the sieve-tube members and companion cells, although the value is essentially the same as between the various parenchymatic elements of the phloem. Compared to several previously studied species, the frequency of plasmodesmata between cell types of the spinach leaf is low. These results are discussed in relation to apoplastic vs. symplastic solute transport and sieve-tube loading in this species.     

Sucrose transport into the phloem of Ricinus communis L. seedlings as measured by the analysis of sieve-tube sap

Careful cutting of the hypocotyl of Ricinus communis L. seedlings led to the exudation of pure sieve-tube sap for 2–3 h. This offered the possibility of testing the phloem-loading system qualitatively and quantitatively by incubating the cotyledons with different solutes of various concentrations to determine whether or not these solutes were loaded into the sieve tubes. The concentration which was achieved by loading and the time course could also be documented. This study concentrated on the loading of sucrose because it is the major naturally translocated sieve-tube compound. The sucrose concentration of sieve-tube sap was approx. 300 mM when the cotyledons were buried in the endosperm. When the cotyledons were excised from the endosperm and incubated in buffer, the sucrose concentration decreased gradually to 80–100 mM. This sucrose level was maintained for several hours by starch breakdown. Incubation of the excised cotyledons in sucrose caused the sucrose concentration in the sieve tubes to rise from 80 to 400 mM, depending on the sucrose concentration in the medium. Thus the sucrose concentration in the sieve tubes could be manipulated over a wide range. The transfer of labelled sucrose to the sieve-tube sap took 10 min; full isotope equilibration was finally reached after 2 h. An increase of K⁺ in the medium or in the sieve tubes did not change the sucrose concentration in the sievetube sap. Similarly the experimentally induced change of sucrose concentration in the sieve tubes did not affect the K⁺ concentration in the exudate. High concentrations of K⁺, however, strongly reduced the flow rate of exudation. Similar results were obtained with Na⁺ (data not shown). The minimum translocation speed in the sieve tubes in vivo was calculated from the growth increment of the seedling to be 1.03 m·h^-1, a value, which on average was also obtained for the exudation system with the endosperm attached. This comparison of the in-vivo rate of phloem transport and the exudation rate from cut hypocotyls indicates that sink control of phloem transport in the seedlings of that particular age was small, if there was any at all, and that the results from the experimental exudation system were probably not falsified by removal of the sink tissues.Abbreviations PTS 3-hydroxy-5,8, 10-pyrenetrisulfonate     

ONTOGENY AND STRUCTURE OF THE SECONDARY PHLOEM IN PYRUS MALUS

Evert , Ray F. (U. Wisconsin, Madison.) Ontogeny and structure of the secondary phloem in Pyrus malus. Amer. Jour. Bot. 50(1): 8–37. Illus. 1963.—The secondary phloem of apple consists of sieve-tube elements, companion cells, phloem parenchyma cells, fiber-sclereids, and ray parenchyma cells. The sieve-tube elements are generally long, slender cells with very oblique end walls and much-compounded sieve plates. All sieve-tube elements initially possess nacreous thickenings. Similar wall thickenings were observed in the differentiating fiber-sclereids and xylem elements. Of the 245 sieve-tube elements critically examined, 242 were associated with companion cells. All of the companion cells were shorter than their associated sieve-tube elements. Young companion cells possess slime bodies which later become dispersed. Callose is often found on the sieve-tube element side of the common wall between sieve-tube element and companion cell. In several collections, callose was found on both sides of that wall. The parenchyma cells are of 3 types: crystal-containing cells; tannin-and/or starch-containing cells; and those with little or no tannins or starch. Any type parenchyma cell may be on to genetically related to a sieve-tube element, that is, may be derived from the same phloem initial as the sieve-tube element. Morphologically, the phloem parenchyma cells intergrade with the companion cells, the tannin- and starch-free parenchyma cells often being difficult to distinguish from companion cells. Most of the tannin- and starch-free parenchyma cells collapse when the contiguous sieve-tube elements become nonfunctional. The fiber-sclereids arise from parenchyma cells which overwinter on the margin of the cambial zone and differentiate in nonfunctional phloem.