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.     

THE ORGANIZATION OF THE VASCULAR SYSTEM IN THE STEMS OF THE NYMPHAEACEAE. I. NYMPHAEA SUBGENERA CASTALIA AND HYDROCALLIS

The vascular system in the stems of Nymphaea odorata and N. mexicana subgenus Castalia, and N. blanda subgenus Hydrocallis consists of continuing axial stem bundles with eight being the usual number. The stem bundles are concentric and xylem maturation is mesarch. Xylem elements consist of tracheids with spirally or weakly reticulated secondary wall thickenings. The phloem is made up of companion cells and short sieve tube members with simple sieve plates that are nearly transverse. At the node each leaf is supplied with two lateral leaf traces and a median leaf trace. A root trace is also present and supplies a series of adventitious roots borne on the leaf base. Flowers and vegetative buds develop directly from the apical meristem and occupy leaf sites in a single genetic spiral. Each flower or vegetative bud is related to a leaf through specific spatial and vascular association. The related leaf is separated from the related flower by three members of the genetic spiral and occupies an adjacent orthostichy. Vascular tissue for the related flower arises from the inner surfaces of the four stem bundles supplying leaf traces to the related leaf and extends through the pith to the flower or vegetative bud via a peduncle fusion bundle. The vascular system organization in the investigated species of Castalia and Hydrocallis is not typically monocotyledonous or dicotyledonous, nor can it be considered transitional between them. The ontogeny of the vascular system is similar to typical dicotyledons and the investigated species of Nymphaea can, therefore, be considered to represent highly specialized and modified dicotyledons.     

VASCULAR ANATOMY OF THE NODAL REGION IN POPULUS DELTOIDES BARTR

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.     

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.     

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.     

VASCULARIZATION OF A MULTILACUNAR SPECIES: POLYSCIAS QUILFOYLEI (ARALIACEAE) I. THE STEM

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.     

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 OCCURRENCE OF “ROOT GAPS” IN THE RHIZOME SOLENOSTELE AND LEAF TRACE OF DENNSTAEDTIA CICUTARIA

The vascular system of the rhizome axes of Dennstaedtia cicutaria consists of a solenostele with an amphicribral vascular bundle centrally located in the pith. Each leaf has a single trace which is an amphicribral vascular bundle. At each node of an axis there is a complex consisting of the main axis, leaf base, and a branch axis attached to the basiscopic margin of the leaf base. Numerous roots are present on the rhizomes and the abaxial side of the leaf bases. Parenchymatous gaps occur in the rhizome solenostele and the leaf trace directly above the departure of some of the root traces. These gaps are termed root gaps. In some instances the root gaps are confluent. However, not all of the root traces have an associated root gap. The leaf trace is inserted laterally on the main and branch axes at the node so that the acroscopic leaf trace margin anastomoses with the main axis of the vascular system and the basiscopic margin with that of the branch axis. Two leaf gaps are associated with each leaf trace, one occurring in the main axis solenostele and the other in the branch axis solenostele. The medullary bundle of each axis anastomoses with each leaf trace at its point of attachment to the rhizome solenostele. Thus, the medullary bundle forms a continuous vascular strand from leaf trace to leaf trace in any given rhizome axis.     

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.     

SHOOT VASCULAR SYSTEM AND PHYLLOTAXIS OF CASUARINA (CASUARINACEAE)

In species of Casuarina with multileaved whorls, each stem vascular bundle divides radially into two at the site of a leaf trace separation, and the same two bundles rejoin acropetally to where the trace supplies a leaf. Such divisions are divisions of a single vascular bundle, and the rejoining of bundles forms a single bundle. Proposals that the extant primary vascular systems of dicotyledons may have been derived as in conifers are incorrect in so far as Casuarina is concerned, or the system has evolved beyond that so far proposed for dicotyledons. Reasons are offered, however, for considering that fernlike leaf gaps are not present. Leaf traces supply leaves at the first nodes distal to their origins. The ways by which an increase or decrease of stem bundles occur are described. Phyllotactic patterns range from helical (rare) to whorled. In the embryo, where leaves occur decussately, of certain species with multileaved whorls, and in the shoot apices of species with tetramerous whorls, slight differences in the levels of leaf attachments and the bending of leaf traces indicate the probable evolution of extant whorled phyllotaxies from one or more helical arrangements. Stages in the evolution are suggested. The leaves in most species with multileaved whorls are in true whorls. The original periderm of branchlets lies internally to the internodal traces and chlorenchyma, but is otherwise external to the vascular system. It is concluded that each leaf originates at its level of separation from the axis despite several structural features suggesting that the leaf bases have become congenitally adnate to the stem.     

VASCULAR SYSTEM OF THE FLORET OF LEERSIA VIRGINICA (GRAMINEAE-ORYZOIDEAE)

The vascular system of the floret of Leersia is unified yet is segmented according to the appendages it serves. The rachilla at the floret base contains a collateral bundle related to the median trace of the lemma. The palea median trace joins the posterior of this bundle in the rachilla as the lemma laterals merge with the anterior. Although the stamen traces enter at the flanks of this rachilla bundle, they do not become fully incorporated into the system until near the floret base where the rachilla bundle, lemma laterals, and palea laterals converge. Traces from the lodicules attach to the anterior of the stamen traces. The base of the vascular system of the pistil, the pistil plexus, attaches tenuously by a bundle to the lower system between the entrance of the stamen traces. A bundle from each style attaches near the anterior of the pistil plexus below the level where the posterior of the pistil plexus rises, as the placental bundle, to merge with the ovule. Characteristics of the vascular system of Leersia, such as the relative discreteness of the staminal and stylar traces and the lack of both the anterior pistil bundle and the xylem discontinuity, are useful for delimiting the Oryzoideae from the Festucoideae.     

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.     

Distinctive tracheid microstructure in stems of Victoria and Euryale (Nymphaeaceae)

Scanning electron microscopy (SEM) photographs of thick sections from liquid‐preserved stems of Victoria cruziana and Euryale ferox show accretions of coarse fibrils on pit membranes of tracheids. The first‐deposited fibrils are randomly orientated; on top of them (facing the tracheid lumina) are axially orientated coarse fibrils. The two systems are interconnected. Axially orientated fibrils were more extensively observed in Euryale than in Victoria and tips of fibrils in Euryale extend over the pit apertures onto secondary wall surfaces. Tracheid–parenchyma interfaces bear rudimentary coarse fibrils on the tracheid side. End walls of Victoria tracheids have highly porose pit membranes, thinner and less complex than those of the lateral intertracheid walls. The structures reported in Victoria and Euryale are consistent with those concurrently reported for stems of other Nymphaeaceae. Although also present in Cabombaceae, the coarse fibrils are otherwise not reported for stems of angiosperms and are not yet reported in roots of any species. Pit membrane remnants in perforation plates of various woody dicotyledons represent a nonhomologous phenomenon. The accretions of coarse fibrils in stem tracheids of Nymphaeaceae do not appear to enhance conduction, although they do contain porosities interconnecting tracheids. Removal of pit membrane remnants from perforation plates of primitive dicotyledon woods by hydrolysis does, on the contrary, suggest conduction enhancement. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 52–57.     

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.     

Floral development in Nymphaea tetragona (Nymphaeaceae)

We describe in detail the floral ontogeny of Nymphaea tetragona from a wild population to provide evidence regarding the phylogenetic position of Nymphaea and to reveal evolutionary trends of flowers in Nymphaeaceae by comparison with that of the other genera. Four sepals are initiated unidirectionally. The basal petals are initiated unidirectionally and alternate with the sepals. The dome‐shaped floral apex continues to expand and produces more petal and stamen primordia. The remaining petals and all stamens are initiated in spirals or whorls. Later, the periphery of the floral apex grows more quickly than the centre and results in a depression in the centre of the apex after all stamens have been initiated. Carpels are simultaneously initiated in a cycle at the periphery of the depression. They are ascidiate. After all organs have been initiated, the centre of the depression on the floral apex grows and develops into a globular structure. The connected inferior ovary, stigma caps and the globular floral apex together form an extragynoecial compitum. Within Nymphaeaceae, the floral ontogeny of Nymphaea is most similar to that of Euryale and Victoria. It differs more from Ondinea and Barclaya, and differs most from Nuphar. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 211–221.     

VASCULAR PATTERNS IN STEMS OF THE CYCLANTHACEAE

A survey was made of the distribution of stem vascular bundles in representatives of ten genera of the tropical monocotyledonous family Cyclanthaceae. Films of series of serial transverse sections were used to reconstruct the stem vasculature. Each leaf trace, followed in a basipetal direction from its level of insertion at the stem periphery, describes an obliquely downward course, initially contacting from 1 to 4 (or more) existing axial bundles. The associated bundles form a compound vascular bundle in which the original bundles initially remain discrete, most commonly in a tetrapolar arrangement, with four separate strands. Followed further in the basipetal direction, the strands eventually fuse partly or completely, usually to form a collateral or amphivasal axial bundle which participates in a new structural cycle. Quantitative variation between different taxa includes a simple pattern in Ludovia, in which only bipolar bundles are developed. More elaborate forms have multipolar bundles with more than four separate strands. A systematically useful observation is that stem vasculature in Cyclanthus, representing the subfamily Cyclanthoideae, does not differ significantly from that in subfamily Carludovicoideae although there are some distinctive structural features.     

INVESTIGATIONS OF NORTH AMERICAN CYCADEOIDS: TRUNKS FROM WYOMING

Delevoryas , Theodore . (University of Illinois, Urbana.) Investigations of North American cycadeoids: trunks from Wyoming . Amer. Jour. Bot. 47(9): 778–786. Illus. 1960.—A study was undertaken of the cycadeoid trunks from Wyoming to more fully understand the morphology and evolutionary significance of these trunks and to determine the validity of the genus Cycadella. The stem named Cycadeoidea (or Cycadella) utopiensis is not the same morphologically as those from Carbon County, Wyoming, and because of its unknown source, it is not a reliable specimen for the study of cycadeoids from the Morrison formation. The Morrison, Carbon County specimens included in Ward's genus Cycadella were shown to have trunk structure, leaf bases, and cones like those in Cycadeoidea. The cone vascular bundle is derived from a fusion of 4 leaf traces. From the fused bundle complex, 4 leaf traces are separated farther out in the cortex, and 1 of these supplies the subtending leaf. In spite of the relatively complex pattern of cone trace formation, there does not seem to be any significant means of distinguishing specimens called Cycadella from the more common Cycadeoidea. Furthermore, it is evident that the genus Cycadeoidea represents a fairly homogeneous group with a wide geographic and geologic range. Pattern of formation of cone traces is interpreted as possible further evidence for regarding cycadeoid cones as portions of foliar systems.     

BIFURCATION OF THE STEM APEX IN ASCLEPIAS SYRIACA

In Asclepias syriaca the overall inflorescence is an anthoclad in which the peduncles are non-axillary, each occurring about 60° away from the axil of a leaf. Ontogenetically, a peduncle is initiated when the stem apex expands laterally and bifurcates into separate apices, neither of which is subtended by any type of organ. One of the two apices continues as the functional apex of the stem (bifurcating again at each subsequent node), and the other functions as the apex of the peduncle. The peduncle first produces a bract and, then, a pedicel in the axil of the bract. Subsequent pedicels are each axillary to separate bracts. The pedicels, therefore, can be interpreted as ordinary lateral branches. However, because the bifurcations of the stem apex are not associated with subtending organs, the branching of the stem does not conform to expected monopodial or sympodial systems in the angiosperms. This suggests the possibility that each bifurcation of the stem apex is a true dichotomy. The anthoclad axis, thus, is a series of dichotomies. Although such a series may have been phylogenetically derived from a monopodial or sympodial ancestor, it is also possible that it may have been retained from a primitive, dichotomizing inflorescence.