VASCULAR ORGANIZATION IN SHOOTS OF CACTACEAE. I. DEVELOPMENT AND MORPHOLOGY OF PRIMARY VASCULATURE IN PERESKIOIDEAE AND OPUNTIOIDEAE

Primary shoot vasculature has been studied for 31 species of Pereskioideae and Opuntioideae from serial transections and stained, decorticated shoot tips. The eustele of all species is interpreted as consisting of sympodia, one for each orthostichy. A sympodium is composed of a vertically continuous axial bundle from which arise leaf- and areole-trace bundles and, in many species, accessory bundles and bridges between axial bundles. Provascular strands for leaf traces and axial bundles are initiated acropetally and continuously within the residual meristem, but differentiation of procambium for areole traces and bridges is delayed until primordia form on axillary buds. The differentiation patterns of primary phloem and xylem are those typically found in other dicotyledons. In all species vascular supply for a leaf is principally derived from only one procambial bundle that arises from axial bundles, whereas traces from two axial bundles supply the axillary bud. Two structural patterns of primary vasculature are found in the species examined. In four species of Pereskia that possess the least specialized wood in the stem, primary vascular systems are open, and leaf traces are mostly multipartite, arising from one axial bundle. In other Pereskioideae and Opuntioideae the vascular systems are closed through a bridge at each node that arises near the base of each leaf, and leaf traces are generally bipartite or single. Vascular systems in Pereskiopsis are relatively simple as compared to the complex vasculature of Opuntia, in which a vascular network is formed at each node by fusion of two sympodia and a leaf trace with areole traces and numerous accessory bundles. Variations in nodal structure correlate well with differences in external shoot morphology. Previous reports that cacti have typical 2-trace, unilacunar nodal structure are probably incorrect. Pereskioideae and Opuntioideae have no additional medullary or cortical systems.     

ONTOGENY OF THE STEM-NODE-LEAF VASCULAR CONTINUUM OF AUSTROBAILEYA

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.     

Developmental features of the discontinuous stem vascular system in the rattan palm Calamus (Arecaceae-Calamoideae-Calamineae)

Calamus is a climbing palm marked by considerable internodal extension and limited stem-thickening growth, but with a surprisingly discontinuous axial vascular system. Stem bundles end blindly in a basipetal direction and are connected to each other only by narrow and late-developing transverse commissures. Vascular connection via leaf traces between stem and leaf is made over about nine plastochrons (P), but the dominant central system is completed by about P(7), with subsequent bundles forming the crowded fibrous peripheral system, which has reduced or no vascular tissues. The stem internode below a leaf completes its extension and maturation only by P(10) to P(11). Axial stem bundles originate as procambial strands that are discontinuous apically for up to 15 plastochrons before being "captured" by a developing leaf. Their distal unconnected ends arise by dedifferentiation of ground parenchyma cells. Protoxylem is initiated as short overlapping initials that differentiate progressively during extension growth, which ruptures all but the last-formed elements. Their form, with tapered ends, means that they mature as tracheids. Metaxylem appears only late in shoot development, shortly before internodal elongation ceases (P(8)) and always unconnected to the late-differentiating protoxylem. In each axial bundle protophloem differentiates as a single strand, subsequently and much later appearing as two separate metaphloem strands as the early initials, ruptured by extension growth, are replaced by fibers. It is suggested that the unique features of this stem can be ascribed to the absence of a "meristematic cap," which otherwise typifies palms of normal habit, and that discontinuity is causally related to the pronounced late stem extension growth.     

VASCULAR PATTERNS IN STEMS OF ARACEAE: SUBFAMILY MONSTEROIDEAE

Analysis of stem vasculature in representatives of subfamily Monsteroideae (Araceae) by cinematographic techniques based on serial sections shows three main patterns of organization. One group of five genera (Rhaphidophora, Epipremnum, Amydrium, Scindapsus, Monstera) is characterized by simple vascular bundles and axial bundles which are derived by basal aggregation of small bundles branching from existing axial bundles. Another group of two genera (Stenospermation, Rhodospatha) is characterized by compound vascular bundles which are made by rather irregular association of individual collateral bundles. These two groups correspond to the tribe Monstereae. The last group which corresponds to the tribe Spathiphylleae includes two genera (Spathiphyllum, Holochlamys) with amphivasal vascular bundles which are highly condensed and irregularly anastomosing. In part, this division is correlated with habit and habitat. Some members of the first group resemble genera within the subfamily Pothoideae quite closely and indicate that the two subfamilies are not clearly circumscribed. Compound bundles in Rhodospatha and Stenospermation do not have the precise organization previously reported for the Pandanaceae.     

FLORAL ANATOMY IN THE SAXIFRAGACEAE SENSU LATO. II. SAXIFRAGOIDEAE AND ITEOIDEAE

The floral anatomy and morphology of 26 species from the Saxifragoideae and three from the Iteoideae are described and compared. The flowers of the Saxifragoideae are predominantly actinomorphic, partially epigynous and/or perigynous, and pentamerous, with two carpels which bear numerous ovules. There is usually some degree of independence between carpels, and the normally separate styles possess both a canal and transmitting tissue. Generally, staminodia are absent and nectariferous tissue, which is not vascularized, is present. The subfamily is characterized by large multicellular trichomes with globular, often glandular, heads. Placentation may be parietal, axile, or transitional between the two; parietal appears to be a derived condition in the subfamily. The vascular cylinder in the pedicel generally consists of several to many discrete bundles from which diverge ten compound traces at the base of the receptacle, leaving an inner cylinder of vascular strands that coalesce at a higher level into either as many ventral bundles as carpels or twice that number. In the former case, each ventral bundle consists of one-half of the vascular supply to each adjacent carpel and separates into individual ventral strands in the distal half of the ovary. The ventral bundles provide vascular traces to the ovules and, along with the dorsals, extend up the style to the stigma. Each trace diverging in a sepal plane typically supplies one or more carpel-wall bundles, a median sepal bundle, and a stamen bundle. Each petal-plane trace usually provides one or more carpel-wall bundles, a lateral trace to each adjacent sepal, a petal bundle and, in flowers with ten stamens, a stamen bundle. Dorsal carpel bundles are usually recognizable and may originate from traces in either perianth plane. While the position of Ribes remains problematical, its floral structure does not easily exclude it from the Saxifragoideae. Floral structure in the Iteoideae is remarkably similar to that in the Saxifragoideae, the main differences being a lesser degree of independence between carpels, generally narrower placentae with somewhat fewer ovules, and the presence of only unicellular, acutely pointed epidermal hairs as opposed to the relatively complex, multicellular trichomes prevalent in the Saxifragoideae.     

COMPARATIVE STUDY OF THE FLORAL VASCULATURE IN SAURURACEAE

The floral vascular systems are compared among all six taxa of Saururaceae, including the two species of Gymnotheca which have not been studied previously. All are zygomorphic (dorsiventrally symmetrical), not radial as sometimes reported, in conformity with dorsiventral symmetry during organogenesis. Apocarpy in the two species of Saururus (with four carpels and six free stamens) is accompanied by a vascular system of four sympodia, each of which supplies a dorsal carpellary bundle, two ventral carpellary bundles, and one or two stamen traces. The level at which the ventral bundles diverge is the major difference in vasculature between the two species. The other four taxa are all syncarpous, and share some degree of stamen adnation and/or connation. The vascular systems also show varying degrees of fusion. The two species of Gymnotheca (with four carpels and six stamens) are very similar to each other; in both, the ventral traces of adjacent carpels fuse to form a placental bundle, which supplies the ovules and then splits into a pair of ventral strands. The flowers of Houttuynia cordata (with only three carpels and three adnate stamens) are sessile. Each flower is vascularized by three sympodia; the median adaxial sympodium is longer than the other two sympodia before it diverges to supply the adaxial organs. Three placental bundles also are formed in Houttuynia, but the three bundles differ in their origin. The median abaxial placental bundle diverges at the same level as the three sympodial bundles of the flower, while the other two lateral placental bundles diverge at a higher level from the median adaxial sympodium. Anemopsis californica, with an inferior ovary of three carpels, sunken in the inflorescence axis, and six stamens adnate to the carpels, has a vascular system very similar to that of Houttuynia cordata. The modular theory of floral evolution is criticized, on the bases of the known behavior of apical meristems and properties of vascular systems. The hypothesis is supported that saururaceous plants may represent a line of angiosperms which diverged very early.     

CAULINE VASCULATURE AND LEAF TRACE PRODUCTION IN MEDULLOSAN PTERIDOSPERMS

Medullosa and Sutcliffia specimens from the Paleozoic of North America and Europe are examined to determine the architecture of the cauline vasculature and mode of leaf trace production. Emphasis is placed on the identification and characterization of protoxylem strands and their relationship to leaf trace production. Organization of the primary xylem varies from a single protostele to a dissected stele composed of two to many more or less independent bundles. In Medullosa the bundles of primary xylem are each surrounded by secondary xylem, forming separate segments of vascular tissue (‘steles’ of previous workers). These vascular segments may divide and fuse at different levels in the stem. A definite number of protoxylem strands occur near the periphery of the primary xylem. The protoxylem strands divide at intervals producing protoxylem to the departing leaf traces. Leaf traces thus formed arise from all the vascular segments in a coordinated and predictable way and pass outward through emission areas in the secondary xylem. This type of cauline vascular architecture is compared to that of other seed plants. The vascular system of Medullosa stems is interpreted as a dissected monostele. Sympodial vascular architecture has apparently evolved from a protostele separately within the medullosan pteridosperms.     

VASCULAR CONSTRUCTION AND DEVELOPMENT IN THE AERIAL STEM OF PRIONIUM (JUNCACEAE)

The aerial stem of Prionium has been studied by motion-picture analysis which permits the reliable tracing of one among hundreds of vascular strands throughout long series of transverse sections. By plotting the path of many bundles in the mature stem, a quantitative, 3-dimensional analysis of their distribution has been made, and by repeating this in the apical region an understanding of vascular development has been achieved. In the mature stem axial continuity is maintained by a vertical bundle which branches from each leaf trace just before this enters the leaf base. Lateral continuity results from bridges which link leaf traces with nearby vertical bundles. Development of the provascular system involves a meristematic cap into which the blind ends of vertical bundles can be followed. Leaf traces are produced continuously in association with developing leaf primordia for a period of over 30 plastochrones; they connect with the vertical bundles in the meristematic cap and so establish the essential vascular configuration which is later reorientated through about 90° by overall growth of the crown. The last bundles to differentiate from the leaf do so outside the meristematic cap and thus fail to make contact with the axial system; they appear in the mature axis as blind-ending cortical bundles. Prionium is only distantly related to palms and its vascular histology is quite different. Nevertheless, the course of vascular bundles and the origin of this pattern in the stem resembles that of a palm. It is suggested that we are examining the fundamental pattern of vascular development in large monocotyledons.     

LEAF VASCULATURE IN BARLEY,HORDEUM VULGARE (POACEAE)

The vascular system of the Hordeum vulgare L. leaf consists of multiple longitudinal strands interconnected by transverse bundles. In any transverse section, the longitudinal strands can be categorized into three bundle types: small, intermediate, and large. Individual longitudinal strands intergrade structurally from one bundle type into another as they descend the leaf. At their distal ends, they have the anatomy of a small bundle. As they descend the leaf, most intergrade into intermediate bundle and then into large bundle types. All strands with large bundle anatomy extend basipetally into the stem. Typically, the other longitudinal strands, which do not intergrade structurally into large bundles, do not enter the sheath, but fuse with other longitudinal strands above the junction of the blade with the sheath. Despite the decrease in number of longitudinal bundles entering the sheath, an increase takes place in the total crosssectional area of sieve tubes and tracheary elements. A linear relationship exists between leaf width and total bundle number in the blade but not in the sheath. Moreover, a linear relationship exists between cross-sectional area of vascular bundles and both total and mean cross-sectional area of tracheary elements and thin-walled sieve tubes.     

MORPHOLOGICAL STUDIES OF THE NYMPHAEACEAE SENSU LATO. XVII. FLORAL ANATOMY OF ONDINEA PURPUREA SUBSPECIES PURPUREA (NYMPHAEACEAE)

Classification and phylogeny of the Nymphaeaceae are unresolved. This study provides floral anatomical data that will assist in elucidating generic interrelationships and systematic relationships to other taxa of angiosperms. The floral anatomy of Ondinea purpurea den Hartog subsp. purpurea has been examined utilizing light microscopy. The peduncle possesses stelar vascular bundle complexes and cortical vascular bundles. Cortical bundles terminate within the peduncle. Each bundle complex consists of 2 collateral bundles on the same radius, the inner bundle inverted; 2 protoxylary lacunae occur yet differ in structure and function. Progressing acropetally, the inner xylary lacunae become discrete mesarch strands surrounded centrifugally by a vascular cylinder formed by divisions and anastomosing of the bundle complexes. Together these become the massive receptacular vascular plexus. The plexus provides collateral traces to the floral organs. Each sepal receives 3 traces that separate from the plexus as 1–3 lateral traces. Petals are absent and no vestigial petal traces have been observed. Distally, the plexus forms several large strands of connate gynoecial and androecial traces termed the principal vascular bundles (PVBs). Ventral veins separate from the PVBs and the latter extend acropetally through the outer ovary wall. Branches of the ventrals and PVBs contribute to septal vascular reticula from which each ovule is supplied by one vascular bundle. Each stamen receives 1 trace from branches of the PVBs. The ventrals and PVBs terminate within the carpellary lobes. A comparative anatomical study is offered that supports the inclusion of Ondinea in the Nymphaeaceae sensu stricto.     

COMPARATIVE STUDIES OF THE NODAL AND VASCULAR ANATOMY IN THE NEOTROPICAL CYATHEACEAE. III. NODAL AND PETIOLE PATTERNS; SUMMARY AND CONCLUSIONS

Comparative studies of the nodal and vascular anatomy in the Cyatheaceae are discussed as they relate to the taxonomy and phylogeny of the family. There is in the Cyatheaceae (excluding Metaxya and Lophosoria) a basic nodal pattern consisting of four major phases of leaf trace separations. Abaxial traces arise from the leaf gap margins, and the last abaxial traces from each side of the gap are larger and undergo numerous divisions. Distally adaxial traces separate from the gap margins, and the last adaxial traces are usually larger and undergo multiple divisions. In addition, medullary bundles frequently become petiole strands of the adaxial arc in the petiole. Rarely, cortical bundles form petiole strands in the abaxial arc in the petiole. Leaf gaps of the squamate genera of the Cyatheaceae are fusiform and possess prominent lateral constrictions which result from medullary bundle fusions and the separation of leaf traces. A characteristic petiole pattern is found in all members of the Cyatheaceae. There is an increase in the complexity of the petiole vascular tissue which results in a gradation from the undivided strand in Metaxya, to the three-parted petiole pattern in Lophosoria, and finally to the much-dissected petiole vascular tissue in the advanced genera. Nodal and vascular anatomy data basically support Tryon's phyletic scheme for the family. The Sphaeropteris-Alsophila-Nephelea line shows certain tendencies toward increased complexity of nodal and vascular anatomy, whereas the Trichipteris-Cyathea-Cnemidaria line shows the same anatomical and morphological characters in a direction of increased simplification or reduction.     

Complexity untwined: The structure and function of cucumber (Cucumis sativus L.) shoot phloem

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.     

Leaf vasculature in Zea mays L.

The vascular system of the Zea mays L. leaf consists of longitudinal strands interconnected by transverse bundles. In any given transverse section the longitudinal strands may be divided into three types of bundle according to size and structure: small, intermediate, large. Virtually all of the longitudinal strands intergrade structurally however, from one bundle type to another as they descend the leaf. For example, all of the strands having large-bundle anatomy appear distally as small bundles, which intergrade into intermediates and then large bundles as they descend the leaf. Only the large bundles and the intermediates that arise midway between them extend basipetally into the sheath and stem. Most of the remaining longitudinal strands of the blade do not enter the sheath but fuse with other strands above and in the region of the blade joint. Despite the marked decrease in number of longitudinal bundles at the base of the blade, both the total and mean cross-sectional areas of sieve tubes and tracheary elements increase as the bundles continuing into the sheath increase in size. Linear relationships exist between leaf width and total bundle number, and between cross-sectional area of vascular bundles and both total and mean cross-sectional areas of sieve tubes and tracheary elements.     

Leaf vasculature in sugarcane (Saccharum officinarum L.)

The vascular system of the leaves of Saccharum officinarum L. is composed in part of a system of longitudinal strands that in any given transverse section may be divided into three types of bundle according to size and structure: small, intermediate, and large. Virtually all of the longitudinal strands intergrade, however, from one type bundle to another. For example, virutually all of the strands having large bundle anatomy appear distally in the blade as small bundles, which intergrade into intermediates and then large bundles as they descend the leaf. These large bundles, together with the intermediates that arise midway between them, extend basipetally into the sheath and stem. Most of the remaining longitudinal strands of the blade do not enter the sheath but fuse with other strands above and in the region of the blade joint. Despite the marked decrease in number of bundles at the base of the blade, both the total and mean cross-sectional areas (measured with a digitizer from electron micrographs) of sieve tubes and tracheary elements increase as the bundles continuing into the sheath increase in size. Linear relationships exist between leaf width and total bundle number, and between cross-sectional area of vascular bundles and both total and mean cross-sectional areas of sieve tubes and tracheary elements.     

Morphometric, AFLP and plastid microsatellite variation in populations of Scalesia divisa and S. incisa (Asteraceae) from the Galápagos Islands

Analysis of the stem vasculature of the American climbing palm Desmoncus reveals structural features differing significantly from the Old World rattan genus Calamus . Desmoncus has a more directly continuous vascular system but nevertheless shows a vessel distribution that makes for high hydraulic resistance in the axial xylem. Desmoncus is like Calamus in having a single very wide metaxylem vessel in each central axial bundle and is also without direct vascular contact between protoxylem and metaxylem tracheary elements. However, in Desmoncus the stem vascular bundle system resembles that in tree palms (as has been described in the model palm Rhapis excelsa ) in having a continuing axial bundle that branches from each outgoing leaf trace together with a large number of bridge connections between leaf traces and peripheral axial bundles. Resistance to axial water transport is, however, evident in the narrowness of the continuing metaxylem elements in the peripheral stem vascular region. Desmoncus has scalariform perforation plates with few thickening bars in the metaxylem vessels, unlike the simple perforation plates found in Calamus . Thus, Desmoncus shows only limited convergence in stem vascular architecture toward the extreme modifications found in Calamus . This is not unexpected since it is clear that the climbing habit evolved independently in the two genera.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 142 , 243−254.     

THE ORGANIZATION OF THE VASCULAR SYSTEM IN THE STEMS OF THE NYMPHAEACEAE. II. NYMPHAEA SUBGENERA ANECPHYA,LOTOS, AND BRACHYCERAS

The anatomy and organization of the stem vascular system was analyzed in representative taxa of Nymphaea (subgenera Anecphya, Lotos, and Brachyceras). The stem vascular system consists of a series of concentric axial stem bundles from which traces to lateral organs depart. At the node each leaf is supplied with a median and two lateral leaf traces. At the same level a root trace supplies vascular tissue to adventitious roots borne on the leaf base. Flowers and vegetative buds occupy leaf sites in the genetic spiral and in the parastichies seen on the stem exterior. Certain leaves have flowers related to them spatially and by vascular association. Flowers (and similarly vegetative buds) are vascularized by a peduncle trace that arises from a peduncle fusion bundle located in the pith. The peduncle fusion bundle is formed by the fusion of vascular tissue derived from axial stem bundles that supply traces to certain leaves. The organization of the vascular system in the investigated taxa of Nymphaea is unique to angiosperms but similar to other subgenera of Nymphaea.     

寄生植物锁阳茎的发育解剖学研究

锁阳茎的初生分生组织由原表皮、基本分生组织以及在基本分生组织中呈波浪式环状排列的原形成层束组成。茎的增粗是由于呈波浪式环状排列的维管束,其“波浪”上下幅度逐渐增大,即从“浪”的基部到“浪”顶端维管束数目由4个逐渐增加到10－12个。维管束数目不断增加是由于：（1）由髓射线薄壁细胞反分化产生分生组织束,分生组织束活动产生新的维管束;（2）维管束中分化出一列或几列薄壁细胞,导致该维管束被分化出的薄壁细胞分成2－3个独立的维管束。     

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.     

VASCULARIZATION OF DEVELOPING LEAVES OF GLEDITSIA TRIACANTHOS L. I. THE NODE,RACHIS, AND RACHILLAE

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.     

THE ORGANIZATION OF THE VASCULAR SYSTEM IN THE STEMS OF THE NYMPHAEACEAE. III. VICTORIA AND EURYALE

Organization of the stem vascular system was analyzed in Victoria species and Euryale ferox. The stem vascular system consists of a number of concentrically-organized continuing axial stem bundles. At the node each leaf is supplied with a root trace, two lateral leaf traces, and a median leaf trace. A peduncle fusion bundle is also present at each node. The peduncle fusion bundle supplies vascular tissue to the median leaf trace and to the peduncle trace. Flowers are nonmedian axillary but have specific vascular, spatial, and developmental relationships to leaves in a manner that resembles the genus Nymphaea. On the basis of the analysis of the stem vascular system, Victoria and Euryale are more similar to each other than to Nymphaea. However, the vascular system in Victoria and Euryale is similar enough to Nymphaea to suggest that Nymphaea, Victoria, and Euryale form a natural taxon of unique angiosperms. The organization of the stem vascular system in Victoria and Euryale is dicotyledonous.