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1.
Microautoradiography was used to follow the translocation pathways of 14C-labeled photosynthate from mature source leaves, through the stem, to immature sink leaves three nodes above. Translocation occurred in specific bundles of the midveins and petioles of both the source and sink leaves and in the interjacent internodes. When each of six major veins in the lamina of an exporting leaf was independently spot-fed 14CO2, label was exported through specific bundles in the petiole associated with that vein. When the whole lamina of a mature source leaf was fed 14CO2, export occurred through all bundles of the lamina, but acropetal export in the stem was confined to bundles serving certain immature sink leaves. Cross-transfer occurred within the stem via phloem bridges. Leaves approaching maturity translocated photosynthate bidirectionally in adjacent subsidiary bundles of the petiole. That is, petiolar bundles serving the lamina apex were exporting unlabeled photosynthate while those serving the lamina base were simultaneously importing labeled photosynthate. The petioles and midveins of maturing leaves were strong sinks for photosynthate, which was diverted from the export front to differentiating structural tissues. The data support the idea of bidirectional transport in adjacent bundles of the petiole and possibly in adjacent sieve tubes within an individual bundle.Abbreviations C central leaf trace - L left leaf trace - LPI leaf plastochron index - R right leaf trace  相似文献   

2.
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.  相似文献   

3.
The ontogeny of the major venation in the lamina of Populus deltoides Bartr. leaves was investigated in relation to the development of original procambial bundles, subsidiary bundles, and their derivatives. Serial sections and clearings were used to show that the midrib region is a composite structure consisting of several independent vascular bundles, each of which eventually diverges into the lamina to become a secondary vein. The sequence of events in the ontogeny of major secondary veins is: (1) an original procambial strand develops acropetally and becomes the precursor of the first vascular bundle of the midrib region of the lamina, (2) ground tissue at the forefront of acropetally developing subsidiary procambial bundles differentiates in a wavelike continuum; meristematic regions precede the acropetally developing procambial bundles, (3) discrete subsidiary bundles differentiate in the meristematic regions as they advance acropetally, (4) subsidiary bundles diverge obliquely in the lamina margin giving rise to the secondary veins in a basipetal fashion, and (5) subsequent differentiation and maturation of the secondary veins occurs within the lamina. The original procambial bundles and first-formed subsidiary bundles become the secondary veins of the uppermost portions of the lamina, the next-formed subsidiary bundles become the secondary veins of the middle portions of the lamina, and the last-formed subsidiary bundles become the secondary veins of the lowermost portion of the lamina.  相似文献   

4.
Development of the Populus leaf is presented as a model system to illustrate the sequence of events that occur during the sink to source transition. A Populus leaf is served by three leaf traces, each of which consists of an original procambial trace bundle that differentiates acropetally and continuously from more mature procambium in the stem and a complement of subsidiary bundles that differentiates bidirectionally from a leaf basal meristem. During development these subsidiary bundles maintain continuity through the meristematic region of the node. The basipetally developing subsidiary bunles form phloem bridges that serve to integrate adjacent leaf traces of the stem vasculature. Distal to the node the acropetally developing bundles from all three leaf traces are reoriented in a precise and orderly sequence to form tiers of petiolar bundles. These tiers of bundles extend into the midrib where bundles diverge at intervals as the major lateral veins. The dorsal-most tier of bundles extends to the lamina tip and each successive tier of bundles contributes to lateral veins situated more proximally in the lamina. Although the midrib and the major vein system differentiate acropetally in the lamina, they mature basipetally. Maturation of the mesophyll and other lamina tissues also mature basipetally. As a consequence of the basi-petal maturation process, the lamina tip matures very early and begins exporting photosynthates while the lamina base is still importing from other leaves. The transition of a leaf from sink to source status must therefore be considered as a progression of structural and functional events that occur in synchrony.  相似文献   

5.
Foster , Adriance S. (U. California, Berkeley), and Howard J. Arnott . Morphology and dichotomous vasculature of the leaf of Kingdonia uniflora. Amer. Jour. Bot. 47 (8): 684–698. Illus. 1960.—An intensive study of the nodal anatomy, petiolar vasculature and open dichotomous venation of the leaf of Kingdonia has revealed a type of foliar vascular system of unusual morphological and phylogenetic interest. The vascular supply at the nodal level consists of 4 collateral traces which diverge from a single gap into the sheathing leaf base. This type of nodal anatomy is perhaps primitive, and comparisons are made with the unilacunar nodes and the 2- and 4-parted leaf trace systems characteristic of many angiospermous cotyledons and the foliage leaves of certain woody ranalian genera. The petiole of Kingdonia is vascularized by 2 pairs of bundles which represent the upward continuation of the 4 leaf traces. A transition from an even (4) to an odd (3) number of strands occurs near the point of attachment of the 5, lobed, cuneiform lamina segments to the petiole. Each of the 2 abaxial bundles dichotomizes and the central derivative branches fuse to form a double bundle which enters the base of the median lamina segment. The 2 adaxial petiolar bundles diverge right and left into the bases of the paired lateral segments of the lamina. An analogous type of transition from an even to an odd number of veins occurs in many angiospermous cotyledons which develop a definable mid-vein. But, in Kingdonia, the bundles which enter the bases of the lamina segments give rise to systems of dichotomizing veinlets devoid of “mid-veins.” Although the majority of the terminal veinlets enter the marginal teeth of the lamina segments, “blind” endings, unrelated to the dentations, occur in all the leaves studied. Typically, all of the vein endings in a given lobule of a lamina segment are derived from the same dichotomous vein system. However, in some leaves, a veinlet dichotomizes directly below a sinus and the branches diverge into the marginal regions of 2 separate lobules. The phylogenetic significance of the occurrence of open dichotomous venation in such an herbaceous angiosperm as Kingdonia is briefly discussed. From a purely morphological viewpoint, the Kingdonia type of venation invites direct comparison with the venation of Sphenophyllum, certain ferns or Ginkgo rather than with any of the known reticulate venation patterns of modern angiosperms. Although the foliar venation of Kingdonia may represent the result of evolutionary reversion, the very rare anastomoses which occur seem primitive in type rather than “vestiges” of a former system of closed venation.  相似文献   

6.
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.  相似文献   

7.
Leaves of Gleditsia triacanthos L. are served by three leaf traces that subdivide in the node to produce subsidiary bundles. The subsidiary bundles differentiate basipetally in the stem and acropetally in the petiole using the original leaf trace bundles (those that developed acropetally) as templates for their development. Within the pulvinus, the acropetal bundle components merge to form the rachis vasculature consisting of a semicircular arc and a ventral chord; several small bundles diverge to form ventral ridge bundles. Mixing of bundles occurs during vascularization of the lateral rachillae axes. Each diverging rachilla axis receives bundles from the semicircular arc, the ventral chord, and a ridge bundle in a relatively reproducible and predictable pattern. During this process the main rachis vasculature is gradually depleted, but the ridge bundles are reconstituted following divergence of each rachilla pair. The distal rachilla pair is vascularized by a bilateral partitioning of the entire rachis vasculature; a remnant of the central leaf trace terminates in a subulate terminal appendage. Vascularization of the bipinnate G. triacanthos leaf is compared to that of the simple Populus deltoides leaf.  相似文献   

8.
Developmental study of the stem-node-leaf vascular continuum of Austrobaileya scandens White reveals that the vasculature within each leaf originates from a single procambial strand, that becomes separated into two strands only at the junction of leaf and stem. At lower levels in the stem the two strands become incorporated into independent portions of the stele. At later stages of development the solitary vascular bundle within the young leaf undergoes considerable lateral growth, resulting in an essentially continuous arc of vascular tissue. Ontogenetic evidence indicates that the vascular bundle in the midrib of the lamina should be regarded as a fundamentally single bundle and not interpreted as two bundles that have undergone various degrees of secondary fusion. A condition of two totally separate bundles extending the entire length of the leaf was not encountered. Our observations confirm the characterization of Austrobaileya as an example of “second rank” level of leaf vasculature. Nodal anatomy emphasizes the extremely isolated taxonomic position of Austrobaileya within the primitive dicotyledons.  相似文献   

9.
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.  相似文献   

10.
Anatomical features of the petiole in several species of Jatropha L. (Euphorbiaceae) are presented as evidence in support of infrageneric relationships. A trilacunar 3-trace nodal pattern is typical for the genus. The vascular supply to the stipules is derived from the branching of the two peripheral leaf traces. The number of vascular bundles range from 11 through 9, 7, 5 and 3, and occur in a ring, as free traces, a medullated cylinder, or as U-shaped free traces. The reduction from nine to three bundles is correlated with the gross morphological features while 11, which occurs only in the section Peltatae (Pax) Dehgan & Webster, presents an increase. Reduction in the number of petiolar traces follows the evolutionary advancement of various taxa. This reduction in traces corresponds with south-north distribution of the species and consequential adaptation to colder and more arid climates in Central America and Africa. Smaller leaves, fewer primary veins and fewer vascular traces have resulted as a response to reduced need for water. Presence of dorsal (super-numerary) bundles which supply the petiolar glands in subgenus Jatropha (= Adenoropium Pax) is considered significant, since African taxa of the section (subsection Pubescentes Pax) have retained these bundles despite the loss of petiolar glands. The latter glands are prominent in the South American and Indian species. Sectional lines in the genus can, therefore, be drawn generally on the basis of numerical constancy and relative uniformity in the arrangement of petiolar traces. The continuity of vascular bundles from the stem into the petiole and variations of bundle arrangements are depicted in three-dimensional drawings.  相似文献   

11.
Summary The ability of a developing cottonwood (Populus deltoides Bartr.) leaf to export 14C-labeled assimilates begins at the lamina tip and progresses basipetally with increasing LPI. This progression indicates that portions of leaves function quasi-independently in their ability to export 14C-photosynthate. Although most of the exported radioactivity was recovered in the petiole as water-80% alcohol-soluble compounds, there was also substantial incorporation into the chloroform and insoluble fractions. This observation indicates that assimilates translocated from the lamina are used in structural development of the petiole. Freeze substitution and epoxy embedding were used to prepare microautoradiographs for localization of water-soluble compounds. Radioactivity was found in all cell types within specific subsidiary bundles of the petiole. However, radioactive assimilates appeared to move from the translocation pathway in the phloem toward active sinks in the walls of the expanding metaxylem cells. Translocation in the mature xylem vessels was not observed.  相似文献   

12.
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.  相似文献   

13.
14.
The nodal anatomy of Xanthophyllum is unilacunar with a single broad trace departing the cauline stele. The "stipular glands" or extra floral nectaries of some species are vascularized by bundles originating from the base of the leaf trace. Considerable variation exists among species in petiole vasculature with siphonosteles, steles with medullary bundles and simple, flat traces present. The lamina also shows variation in the presence or absence of a hypodermis, nature of vein sheathing, presence or absence of abaxial epidermal papillae, amount of intercellular spaces, and mature stomatal patterns which range from anisocytic and paracytic to those in which no subsidiary cells are discernible. Of nearly uniform occurrence throughout the genus are extraordinary tracheoid foliar idioblasts, which are confined to the veins in terminal or subterminal positions. The large amount of variation in leaf anatomy is shown to be taxonomically significant within the genus.  相似文献   

15.
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.  相似文献   

16.
Cucurbit phloem is complex, with large sieve tubes on both sides of the xylem (bicollateral phloem), and extrafascicular elements that form an intricate web linking the rest of the vasculature. Little is known of the physical interconnections between these networks or their functional specialization, largely because the extrafascicular phloem strands branch and turn at irregular angles. Here, export in the phloem from specific regions of the lamina of cucumber (Cucumis sativus L.) was mapped using carboxyfluorescein and 14C as mobile tracers. We also mapped vascular architecture by conventional microscopy and X-ray computed tomography using optimized whole-tissue staining procedures. Differential gene expression in the internal (IP) and external phloem (EP) was analyzed by laser-capture microdissection followed by RNA-sequencing. The vascular bundles of the lamina form a nexus at the petiole junction, emerging in a predictable pattern, each bundle conducting photoassimilate from a specific region of the blade. The vascular bundles of the stem interconnect at the node, facilitating lateral transport around the stem. Elements of the extrafascicular phloem traverse the stem and petiole obliquely, joining the IP and EP of adjacent bundles. Using pairwise comparisons and weighted gene coexpression network analysis, we found differences in gene expression patterns between the petiole and stem and between IP and EP, and we identified hub genes of tissue-specific modules. Genes related to transport were expressed primarily in the EP while those involved in cell differentiation and development as well as amino acid transport and metabolism were expressed mainly in the IP.  相似文献   

17.
Klaus Schmitz 《Planta》1970,92(3):208-221
Summary The petiole of Pelargonium zonale is traversed by 17 bundles, whose arrangement and form are typical for this plant. The bundles of the petiole are connected with the conducting system of the axis and with the main nerves by a system of phloem anastomoses in the leaf base and in the junction between the petiole and the leaf blade (Fig. 2). The anatomical findings were confirmed and extended by a study of the translocation of K-fluorescein and 14C. It could be shown that the metaphloem of the central petiole bundle is composed of phloem subunits, each of which is connected with the phloem of one certain main nerve only (Fig. 4). Accordingly, if fluorescein or 14CO2 is applied to one main nerve, the dye or 14C-material is translocated exclusively in a small phloem area of the central bundle. Autoradiograms of the petioles indicate that the 14C-labelled assimilates (sucrose, glucose, fructose and amino acids) are translocated exclusively in the phloem. A lateral movement of the labelled material within the petiole was not observed. The metaphloem of the central petiole bundle of Pelargonium zonale revealed a functional organization of phloem subunits.

Teil einer Dissertation unter der wissenschaftlichen Leitung von Prof. Dr. J. Willenbrink.  相似文献   

18.
The ontogeny of vascular bundles in the nodal region of Populus deltoides Bartr. was examined to understand more thoroughly the structure-function relation between leaf and stem. Three vascular traces from the stem independently enter each leaf in the nodal region. At the base of each developing leaf a region was observed in which both bundle size and vascular development was reduced; this region was referred to as the constricted zone. The constricted zone was described quantitatively at 13 locations within the nodal region of a leaf at LPI 5 by determining the number of metaxylem vessels and the total metaxylem vessel area in each of the three leaf traces. A plot of these data showed a distinct minimum value for total metaxylem vessel area within the constricted zone of each trace; the location of this minimum value was referred to as the constriction plane. Each vascular bundle within the nodal region is composed of independent subsidiary bundles that originate within the constricted zone. These bundles provide a direct connection between the leaf lamina and the stem. The node was defined anatomically on the basis of the ontogenetic development of the subsidiary bundles. The node began at the initial exit point of the central trace from the vascular cylinder and extended distally to the constriction plane. This definition allowed us to quantify the limits of each node. The origin of the initiating layer and metacambium was also examined within the nodal region. These precursors of the cambium develop continuously and acropetally from the stem into the leaf. The developmental implications of the constricted zone and the metacambium within the nodal region are discussed with respect to wood formation.  相似文献   

19.
The odd-pinnate leaves of Polyscias quilfoylei have a sheathing leaf base that completely encircles the stem. At each node, many traces depart the vascular cylinder and traverse an obliquely upward course through the leaf base before aggregating in the rachis. Lateral traces diverge from parent traces in the stem vasculature at variable times relative to the leaf they serve, from variable positions in the vascular cylinder and from parent traces of variable ages. The stem vasculature is formed by the coalescing of leaf traces from as many as five leaves. All bundles departing the vascular cylinder at a node to serve a leaf are true leaf traces originating independently in the stem. Leaf traces develop acropetally from their positions of origin on parent traces. Primordial leaves are first served by the median trace and later by lateral traces. Many traces were recognized in the internodes subtending embryonic leaves, but they could not be related either to a specific leaf or to a specific position within a leaf. Because these traces had not yet achieved contact with a primordial leaf site, they were assumed to be in the process of developing acropetally at the time of sampling. Observations suggest that the multiple traces in this species might perform a similar function of integrating the vascular cylinder that subsidiary bundles perform in certain uni- and trilacunar species.  相似文献   

20.
We conducted a comparative anatomical study of the petiole of 16 taxa belonging to the tribe Sorbarieae (Rosaceae) (Adenostoma, 2 spp.; Chamaebatiaria, 1 sp.; Sorbaria, 6 spp., 3 vars., and 1 forma; and Spiraeanthus, 1 sp.) and the related genus Lyonothamnus (1 sp. and 1 ssp.). The distal, medial and proximal regions of petioles were transversely sectioned using conventional embedding and staining methods. Cuticles, crystals, trichomes and pericyclic fiber patterns were observed and studied. Three types of vascular nodal patterns were recognized: Type 1 was seen in Chamaebatiaria, Lyonothamnus, and Spiraeanthus (simple‐trace nodal pattern with slightly curved or U‐shaped vascular bundle); type 2 was found in Adenostoma (multiple‐traces nodal pattern with free vascular bundles); and type 3 was unique to Sorbaria (bundles fused to form a siphonostele nodal pattern). Some petiolar anatomical characteristics (e.g. cuticles, crystals, trichomes, vascular nodal pattern, and pericyclic fiber patterns) were found to provide useful information for taxonomic studies within Sorbarieae. On the basis of these characteristics, a dichotomous key for identification at the generic/specific level is provided. We also report a structural change in the vascular bundles from the stem‐leaf transitional zone to the leaf medial zone.  相似文献   

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