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.     

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.     

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.     

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.     

Photosynthetic Characteristics of Portulaca grandiflora, a Succulent C(4) Dicot : CELLULAR COMPARTMENTATION OF ENZYMES AND ACID METABOLISM

The succulent, cylindrical leaves of the C₄ dicot Portulaca grandiflora possess three distinct green cell types: bundle sheath cells (BSC) in radial arrangement around the vascular bundles; mesophyll cells (MC) in an outer layer adjacent to the BSC; and water storage cells (WSC) in the leaf center. Unlike typical Kranz leaf anatomy, the MC do not surround the bundle sheath tissue but occur only in the area between the bundle sheath and the epidermis. Intercellular localization of photosynthetic enzymes was characterized using protoplasts isolated enzymatically from all three green cell types.     

杨树叶薄层培养中不定芽形态发生的细胞组织学研究

将杂种杨树(Populus nigra var.betulifolia×P.trichocarpe)NE299叶主脉用振动切片机横切成400μm或800μm的薄切片,培养在附加0.2mg/L BA和0.01mg/L NAA的木本植物培养基上。培养后,位于主脉维管束两侧中上部的维管束鞘薄壁细胞首先启动分裂。几乎同时,与其邻接的一些栅栏组织细胞也分裂,并很快形成胚性分生细胞团。主脉的愈伤组织主要由维管束鞘薄壁细胞,以及与其邻接的一些栅栏组织细胞和韧皮部的薄壁细胞分裂而来。不定芽通常发生在愈伤组织的周边区,也可以起源于维管组织结节(vascular nodules)周围的形成层状细胞。侧脉的维管束鞘细胞分裂活动很强,可不经愈伤组织直接长成不定芽。杨树叶主脉处的维管束鞘薄壁细胞在与叶肉组织相邻接的细胞中,通常含有少量较小的叶绿体,而位于背腹面的细胞中含有贮藏的淀粉。对形态发生的特定部位及其细胞进行了讨论。     

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.     

虉草营养器官解剖结构与抗旱耐涝性及利用之间的关系

为了从显微结构上进一步探讨虉草(Phalaris arundinacea L.)的抗旱耐涝性及与利用的关系,于2011年采用常规石蜡切片技术,对其根、茎叶3种营养器官进行解剖观察。结果表明,虉草根的结构自外而内依次为表皮、皮层、维管束鞘、初生韧皮部和初生木质部;茎由表皮、基本组织和维管束构成;叶片内部结构可分为表皮、叶肉和叶脉3部分。根皮层大的细胞间隙和气腔,初生木质部的后生大导管和茎基本组织解体形成的髓腔都是虉草良好的通气组织,是其耐水淹的主要显微特征。茎、叶片角质化的表皮和叶表皮所含的丰富泡状细胞组是虉草具有抗旱性的主要解剖结构特征。叶肉细胞排列紧密且只有少量气孔分布于叶片下表皮,这样的结构可减少蒸腾;叶肉细胞富含叶绿体,增强光合作用,获得更多的同化产物,确保了植株在干旱条件下也有足够的光合产物来维持正常的生理活动。茎、叶维管束部分大量的木纤维起到支撑作用。虉草根的皮层和维管柱部分、茎的基本组织和维管束部分、叶的叶脉部分都含有大面积的厚壁细胞,厚壁细胞中含有丰富的粗纤维和木质素。丰富的粗纤维、木质素等成分则是虉草能成为新能源燃料植物的必备条件。     

Chlorophyll a Fluorescence Transients in Mesophyll and Guard Cells : MODULATION OF GUARD CELL PHOTOPHOSPHORYLATION BY CO(2)

Chlorophyll fluorescence transients from mesophyll and guard cell chloroplasts of variegated leaves from Chlorophytum comosum were compared using high resolution fluorescence spectroscopy. Like their mesophyll counterparts, guard cell chloroplasts showed the OPS fluorescence transient indicating the operation of the linear electron transport and the possible generation of NADPH in these organelles. They also showed a slow fluorescence yield decrease, equivalent to the MT transition in mesophyll, suggesting the formation of the high energy state and photophosphorylation. Unlike the mesophyll chloroplasts, the fluorescence from guard cell chloroplasts lacked the increment of the SM transition, indicating that the two types of chloroplasts have some metabolic differences. The presence of CO₂ (supplied as bicarbonate, pH 6.7) specifically inhibited the MT-equivalent transition while its absence accelerated it. These observations constitute the first specific evidence of a guard cell chloroplast response to CO₂. Control of photosynthetic ATP levels in the guard cell cytoplasm by CO₂ may provide a mechanism regulating the availability of high energy equivalents at the guard cell plasmalemma, thus affecting stomatal opening.     

ORGANIZATION AND ONTOGENY OF THE VASCULAR SYSTEM IN THE PETIOLE OF EASTERN COTTONWOOD

The topologic arrangement of petiolar bundles varies within the length of the cottonwood petiole. Each petiolar bundle is formed by the subdivision and aggregation of acropetally differentiating subsidiary bundles in a predictable pattern. The subsidiary bundles provide vascular continuity between the stem and specific portions of the leaf lamina. Spot-labeling of individual veins with ¹⁴CO₂, freeze substitution, and microautoradiography were used to establish the relation between the secondary veins of the lamina and the vasculature of the petiole. Within the petiole vasculature each subsidiary bundle was continuous with a specific portion of the lamina and seemed to have a separate function. Subsidiary bundles continuous with the central leaf trace were closely related functionally to the tip region of the lamina, while the subsidiary bundles continuous with the lateral leaf traces were functionally related to the middle and basal portions of the lamina.     

STUDIES ON THE LEAF OF AMARANTHUS RETROFLEXUS (AMARANTHACEAE): QUANTITATIVE ASPECTS,AND SOLUTE CONCENTRATION IN THE PHLOEM

The vascular system of the leaf of Amaranthus retroflexus L. was examined quantitatively, and plasmolytic studies were carried out on it to determine the solute concentration in cells of the phloem at various locations in the leaf. The proportion of phloem occupied by sieve tubes varies considerably with vein size and leaf size. Collectively, the cross-sectional area of sieve tubes of all tributaries at their points of entry into either a secondary or midvein far exceeds the total cross-sectional area of sieve tubes at the bases of those major veins. In addition, the total volume of sieve tubes in the “catchment area” of a secondary vein is much greater than total sieve-tube volume of the secondary vein itself. The plasmolytic studies revealed the presence of positive concentration gradients in the sieve tubes of the lamina from the minor veins and tips of the secondaries to the bases of the secondaries and from the tip to the base of the midvein. The C₅₀ (the estimated mannitol concentration plasmolyzing, on the average, 50% of the sieve-tube members) was 1.5 m for minor veins and tips of secondary veins and 1.1 m for the bases of secondaries; 1.3 m for the tip of the midvein and 0.6-0.7 m for the midvein in the basal third of the lamina.     

Translocation pathways in the petioles and stem between source and sink leaves of Populus deltoides Bartr. ex Marsh.

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     

Sucrose export defective1 encodes a novel protein implicated in chloroplast-to-nucleus signaling

The Sucrose export defective1 (Sxd1) gene of maize was cloned and shown to encode a novel protein conserved between plants and cyanobacteria. The structure of the Sxd1 locus was determined in wild-type plants and two independent sxd1 alleles. Expression analysis demonstrated that the gene was transcribed in all green tissues, with highest levels in maturing leaf blades. In situ hybridization studies revealed high levels of Sxd1 mRNA in bundle sheath cells, with lower levels within the mesophyll. The SXD1 protein was localized to chloroplasts, in both bundle sheath and mesophyll cells. Levels of sucrose, glucose, and fructose were compared between wild-type and sxd1 plants. Mutant plants were fully capable of producing sucrose and accumulated all three sugars at concentrations above those measured in wild-type plants. Despite these increased sugar concentrations, photosynthetic gene expression was not significantly downregulated in affected areas of sxd1 leaf blades. These results are consistent with photosynthate being trapped within anthocyanin-accumulating regions of sxd1 leaves due to plasmodesmal occlusion at the bundle sheath-vascular parenchyma boundary of the minor veins. A model for SXD1 function is proposed in which the protein is involved in a chloroplast-to-nucleus signaling pathway necessary for proper late-stage differentiation of maize bundle sheath cells, including the developmentally regulated modification of plasmodesmata.     

两种苦丁茶代用品叶结构和后含物与正品的异同

大叶冬青、枸骨叶的结构与苦丁茶基本相似:背腹叶;栅栏组织数层;外韧维管束;小维管束具维管束鞘;气孔仅分布于下表皮,有无规则型和轮列型两种;叶内有纤维、石细胞。叶内都含有单宁、淀粉、脂类及少量草酸钙结晶。叶结构主要区别在于苦丁茶叶主脉维管束成心形的环,另两种呈弧形;枸骨叶缘有横跨叶断面的大纤维束,而另两种叶缘仅少量纤维。     

Transitions in photosynthetic parameters of midvein and interveinal regions of leaves and their importance during leaf growth and development

Abstract: The areal development of photosynthetic efficiency and growth patterns in expanding leaves of two different dicotyledonous species - Coccoloba uvifera and Sanchezia nobilis - was investigated by imaging both processes repeatedly over 32 days. Measurements were performed using combined imaging systems for chlorophyll fluorescence and growth, with the same spatial resolution. Significant differences in potential quantum yield of photosynthesis (F_v/F_m), a parameter indicating the functional status of photosystem II, were found between midvein and interveinal tissue. Although base-tip gradients and spatial patchiness were observed in the distribution of relative growth rate, neither midvein nor interveinal tissue showed such patterns in F_v/F_m. In young leaves, F_v/F_m of the midvein was higher than F_v/F_m of interveinal tissue. This difference declined gradually with time, and upon cessation of growth, F_v/F_m of interveinal regions exceeded those of midvein tissue. Images of chlorophyll fluorescence quenching showed that ΔF/F_m' in the different tissues correlated with F_v/F_m, indicating that, in these uniformly illuminated leaves, transitions in photosynthetic electron transport activity follow those of predawn quantum efficiency. We explore the implications of these observations during leaf development, discuss effects of sucrose delivery from veins to interveinal areas on relative rates of photosynthetic development in these tissues, and propose that the initially higher photosynthetic activity in the midvein compared to the intervein tissues may supply carbohydrates and energy for leaf growth processes.     

Translocation of 14Carbon Within and Between Leaves

Plants of tobacco and a variegated variety of Pelargonium wereused to investigate some aspects of translocation of ¹⁴Carbonwithin and between leaves, following assimilation of ¹⁴CO₂ byone of the leaves. Bi-directional transport in leaves is consideredto result from import into immature regions and exports frommature regions. In variegated leaves the chlorotic areas behavelike immature areas in the sense that they continue to importtranslocate from outside the leaf to a greater extent than adjacentgreen areas. However, some transport occurs from green to chloroticareas via the veins in the same leaf. Using masking techniqueson tobacco leaves it was shown that labelled carbon failed tomove across darkened mesophyll. This was taken to indicate thatthe mechanism resulting in translocation from leaves was locatedin the veins. Labelled carbon was shown to leave the veins ofimporting leaves along their entire length. A simplified technique for freeze-drying plant material priorto wax-embedding is described and some of the limitations ofheat-drying as a preliminary to gross autoradiography of leavesare discussed.     

INTRAMARGINAL VEINED LAURACEAE LEAVES FROM THE ALBIAN–CENOMANIAN OF CHARENTE-MARITIME (WESTERN FRANCE)

Abstract:  Eucalyptolaurus depreii gen. et sp. nov. is proposed for angiosperm leaves newly collected from uppermost Albian – lowermost Cenomanian of Charente-Maritime (western France). They consist of simple, narrow, elongate laminas with entire margins and intramarginal veins. The epidermal cells of adaxial cuticle shows small, rounded, blunt papillae outward that protrude inward and fuse together as rolls along and parallel to the margins, while the adaxial cuticle bears brachyparacytic stomatal apparatus that exhibit sunken guard cells and hair bases consisting of a thick-walled pore surrounded by radially arranged differentiated cells. Resin bodies occur inside the mesophyll. These characters closely resemble the lauroid taxa ' Myrtophyllum ' and Pandemophyllum from the Cenomanian of the Czech Republic and Dakota (USA) respectively. The narrow angle of basilaminar secondaries and the whole suite of features in the guard cells (sunken guard cells embedded into subsidiary cells and stomatal ledges) strongly support close affinity with the Lauraceae. From the Cenomanian lauraceous reproductive organs and their related leaves already showed high disparity and diversity. In addition they displayed a broad ecological range from freshwater floodplains to brackish swamps. This combined to high diversity of reproductive organs suggest ecological radiation of Lauraceae by the Cenomanian.     

Duabanga-like leaves from the Middle Eocene Princeton chert and comparative leaf histology of Lythraceae sensu lato

A permineralized lythraceous leaf type found in close association with fruits, stems, and roots of Decodon allenbyensis Cevallos-Ferriz et Stockey in the Middle Eocene Princeton chert of British Columbia, Canada, is described. Midribs have a prominent C-shaped midvein surrounded by sclerenchyma, with an adaxial epidermis of rectangular to rounded cells lacking enlarged mucilage cells. Leaves are dorsiventral, 180-270 μm thick at the lamina, with a double palisade layer. Abaxial epidermal cells have prominent papillae, and these epidermal cells can be infected by fungi, forming dark sterile stromata. Fossil leaves are similar to those of Myrtales and are compared to those of Lythraceae sensu lato. Although these leaves are thought to belong to the previously described Decodon allenbyensis found in the same chert layer, they lack the diagnostic features of extant Decodon leaves. Instead they share most anatomical similarities with Duabanga grandiflora Roxburgh ex DC Walpers (Lythraceae, subfamily Duabangoideae) including vascular tissues, palisade and spongy mesophyll, bundle fibers, and abaxial epidermal papillae. Duabanga grandiflora differs from the fossil in having mucilaginous cells and a consistently V-shaped abaxial midrib. Although anatomically similar to Duabanga, the fossil leaves are considered those of D. allenbyensis, based on association and the depositional environment prior to preservation. Recent phylogenetic analyses place Duabanga and Decodon in separate clades within Lythraceae, but relationships between these clades are not well supported, indicating that fossil leaves should provide useful anatomical characters for elucidating relationships within Lythraceae.     

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。     

Sink to Source Transition of Populus Leaves

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.