Anatomy of stem‐node‐leaf continuum in Aconitum (Ranunculaceae) in the Eastern Carpathians

Studies of the anatomical structure of the stem and leaf, with special emphasis on the organization of the vascular system, has been carried out on 13 Aconitum species from Aconitum subgenera Aconitum, Anthora and Lycoctonum. All investigated species show a more or less mesomorphic anatomical structure, typical for other Ranunculaceae. Hence, these species have similar trilacunar three‐trace organization of the nodal vascular system. In the stem the vascular system is open (with weakly developed cambium) or closed collateral, and incomplete (consisting of the bundles were represented), large complete, middle‐size complete and small incomplete or with weakly developed xylem. The number of vascular bundles in petioles appear to have no taxonomical value. Nevertheless, it was found that the spatial organization of these vascular bundles in the petiole are of taxonomic importance. As a result, the investigated species can be divided into four main groups congruent with the current sectional and subgeneric division of the genus. The only exception was A. × cammarum (A. sect. Acomarum) in which the is identical to that of A. sect. Cammarum. The most primitive vascularization is found in A. anthora, while the most advanced one is found in A. variegatum. The highly differentiated and distinct nodal anatomy of A. anthora suggests a high, plausibly subgeneric, taxonomical rank of this species.     

Anatomy,trichome morphology and palynology of Salvia chrysophylla Stapf (Lamiaceae)

The anatomy, palynology, morphology and distribution of the trichomes on the aerial parts of Salvia chrysophylla Stapf, an endemic species in Turkey, were studied in order to understand the usefulness of these characteristics for systematic purposes. Some anatomical characters such as (1–)2–24-rowed pith rays in roots, dorsiventral leaves, obviously larger upper epidermal cells, and two to three large vascular bundles in the center and two to four small subsidiary bundles in the wings of petiole provide information of taxonomical significance. Three main types of trichomes were observed on the stem, inflorescence axis, leaf and calyx surfaces of S. chrysophylla. They are peltate, capitate glandular and non-glandular. Capitate glandular and non-glandular trichomes were further subdivided into several kinds. Glandular trichomes are present in abundance on the inflorescence axis and calyx, but non-glandular ones were mainly situated on the leaf and stem. Scanning Electron Microscopy (SEM) studies on the pollen grains have revealed that they are oblate-spheroidal and their exine ornamentation is bireticulate-perforate.     

The anatomical and micromorphological characteristics of <Emphasis Type="Italic">Salvia vermifolia</Emphasis> (Section <Emphasis Type="Italic">Aethiopis</Emphasis> Bentham,Lamiaceae) from Central Anatolia,Turkey

The gross anatomical features, namely stem, blade and petiole and the micromorphological features, trichome, pollen and nutlet of Salvia vermifolia Hedge & Hub.-Mor. (Sect. Aethiopis Bentham, Lamiaceae) were examined using light microscopy (LM) and scanning electron microscopy (SEM). S. vermifolia is a perennial endemic herb growing on igneous and serpentine slopes in Sivas province, Central Anatolia, Turkey. The results obtained from anatomical studies show that the stem is made up of 4–8-layers of collenchyma cells and 1-3-layers chlorenchyma cells. The blade is amphistomatic, bifacial, with 2-3-layered palisade cells on the adaxial. The petiole has three large central bundles and six small subsidiary bundles. Peltate glandular, capitate glandular and non-glandular trichomes are present in S. vermifolia. The peltate glandular trichomes are present in abundance on the calyx and corolla, and capitate glandular trichomes are predominant on the calyx, inflorescence axis, pedicel and stem. The acicular non-glandular trichomes are mainly situated on the calyx, corolla, inflorescence axis and pedicels whereas the floccose non-glandular trichomes are common on both sides of the leaf, inflorescence axis and petiole. The pollen grains are hexacolpate to octacolpate, medium to large (P, polar axis=45.29–62.49 μm; E, equatorial axis=45.32-62.38 μm), oblate-spheroidal to prolate-spheroidal (P/E=0.91–1.02) and their exine ornamentation is bireticulate. The nutlets are 2.70–2.90 mm long and 1.90–2.13 mm wide, brown, ovoid-oblong in their outline, rounded-trigonous in transverse section, and their surface is glabrous and colliculate.     

绞股蓝营养器官的结构及其人参皂甙的组织化学定位研究

绞股蓝是多年生草质藤本植物。根系由不定根组成,根的初生结构木质部为2－4原型,次生结构中栓内层较厚,攀缘茎,具5棱,周围纤维连成一环,幼茎的维管束排成两圈,外圈5个,内圈4或5个,老茎圆柱形,周围纤维呈不连续环状,维管束具次生木质部和次生韧皮部,排成一圈,掌状复叶互生,小叶5－7片,背腹型,叶柄具5束维管束,进入小叶时分为7－9束,茎和叶的初生维管束为双韧维管束,组织化学实验表明,绞股蓝人参皂甙主要分布在营养器官的同化组织及韧皮部薄壁细胞中,厚角组织,表皮及周皮的栓内层也有少量分布。     

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.     

COMPARATIVE ANATOMY OF THE PETIOLE AND INFRAGENERIC RELATIONSHIPS IN JATROPHA (EUPHORBIACEAE)

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.     

INTERNAL PHLOEM IN THE PULVINUS OF SOYBEAN PLANTS

Glycine max, like many species of Fabaceae, has pulvini at the base of the petiole. In this structure, the vascular cylinder is constricted and consists of a ring of phloem surrounding a ring of xylem. A combination of light and transmission electron microscopy and histochemistry showed that, in addition, there are groups of internal phloem strands in the pulvinar pith. This was confirmed by direct observation of sieve plates and crystalline P-protein inclusions typical of leguminous sievetube members. Serial sections through the stem–pulvinus–petiole revealed that a spatial reorientation of the vascular tissue in the pulvinus resulted in the formation of internal phloem strands, which are continuous with the external phloem bundles above and below the pulvinus. Using 6(5)carboxyfluorescein (6CF) as a fluorescent tracer of phloem transport, we have shown that the internal phloem was active. In most of the experiments, when 6CF was applied to a source leaf, the internal phloem was not stained when the stem was girdled between the source leaf and the roots. Thus, we suggest that the internal phloem of the pulvinus of soybean is specialized for transport toward the root.     

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.     

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.     

The role of phytochrome B,D and E in thermoperiodic responses of <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis>

The objective of this work was to study the role of the phytochromes (phy) B, D and E in the thermoperiodic control of elongation and flowering time in Arabidopsis thaliana. WT, and phyB, phyD and phyE single mutants, and phyB phyD and phyB phyE double mutants, were grown under day/night temperatures (DT/NT) of 12/22°C, 17/17°C or 22/12°C (negative, zero and positive DIF, respectively) for inflorescence stem length measurements, and under DT/NT 17/25°C or 25/17°C (negative and positive DIF, respectively) for leaf morphology and flowering time measurements. In WT final length of the stem, petiole and leaf blade were longer under positive DIF compared to negative DIF. The temperature effect was stronger in the leaf petiole than the stem, whereas only a slight change was seen in the leaf blade length direction and none in the width direction. The temperature effect on stem and petiole elongation was reduced or nearly eliminated in the genotypes lacking phyB, while a phyD or a phyE mutation had no influence or a slightly positive influence on the temperature effect, respectively. These results suggest that phyB, and not phyD or phyE, is needed for a complete thermoperiodic control of elongation growth in A. thaliana. For all genotypes tested, plants flowered earlier at negative DIF than positive DIF, suggesting that none of the three phytochromes B, D, or E is needed for a thermoperiodic control of flowering time in A. thaliana.     

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.     

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.     

SECRETORY RESERVOIRS (DUCTS) OF TWO KINDS IN GIANT RAGWEED (AMBROSIA TRIFIDA; ASTERACEAE)

Two types of tubular secretory reservoirs occur in Ambrosia trifida, the first such example known in plants. Paraffin and resin sections, and clearings showed that, although each type consists of many separate unbranched tubes, they differ in anatomy, secretory contents, distribution, and length. Reservoirs (PAR) containing a red substance (presumably a polyacetylene) and lined with a biseriate epithelium parallel the largest leaf and stem vascular bundles. One PAR arises near the base of each leaf lobe midrib and extends through the petiole to the node or continues in the stem cortex to the node below. Other PARs start at the cotyledonary node or in cotyledons and extend down into the primary root, where they have only a single layer of unspecialized epithelium. PARs realign themselves, and more form de novo, until the primary root has two to four separate arrays of PARs abutting the endodermis, each with three to six parallel PARs. Branch roots have similar PAR arrays but unconnected to PARs of the parent root. Inflorescence PARs occur only in bracts, and in petals of male flowers. The second type of reservoir (OR) has a uniseriate epithelium and contains an unidentified oil. ORs occur in phloem, and in pith next to xylem, of stem and large leaf bundles. They dwindle in successively smaller veins until the two smallest orders lack them. ORs occur only in phloem in the hypocotyl; none occur in cotyledons, roots, or floral parts.     

Anatomical changes induced by increasing NaCl salinity in three fodder shrubs, <Emphasis Type="Italic">Nitraria retusa</Emphasis>, <Emphasis Type="Italic">Atriplex halimus</Emphasis> and <Emphasis Type="Italic">Medicago arborea</Emphasis>

Nitraria retusa and Atriplex halimus (xero-halophytes) plants were grown in the range 0–800 mM NaCl while Medicago arborea (glycophyte) in 0–300 mM NaCl. Plants were harvested after 120 days of salt-treatment. The present study was designed to study the effect of salinity on root, stem and leaf anatomy, water relationship, and plant growth in greenhouse conditions. Salinity induced anatomical changes in the roots, stems and leaves. The cuticle and epidermis of N. retusa and A. halimus stems were unaffected by salinity. However, root anatomical parameters (root cross section area, cortex thickness and stele to root area ratio), and stem anatomical parameters (stem cross section area and cortex area) were promoted at 100–200 mM NaCl. Indicating that low to moderate salinity had a stimulating effect on root and stem growth of these xero-halophytic species. At higher salinities, root and stem structures were altered significantly, and their percentages of reduction were higher in A. halimus than in N. retusa whereas, in M. arborea, they were strongly altered as salinity rose. NaCl (100–300 mM) reduced leaf water content by 21.2–56.2% and specific leaf area by 51–88.1%, while increased leaf anatomical parameters in M. arborea (e.g. increased thickness of upper and lower epidermis, palisade and spongy mesophyll, entire lamina, and increased palisade to spongy mesophyll ratio). Similar results were evidenced in A. halimus leaves with salinity exceeding 100 mM NaCl. Leaves of N. retusa were thinner in salt-stressed plants while epidermis thickness and water content was unaffected by salinity. The size of xylem vessel was unchanged under salinity in the leaf’s main vein of the three species while we have increased number in M. arborea leaf main vein in the range of 200–300 mM NaCl. A longer distance between leaf vascular bundle, a reduced size and increased number of xylem vessel especially in stem than in root vascular system was evidenced in M. arborea treated plants and only at (400–800 mM) in the xero-halophytic species. The effects of NaCl toxicity on leaf, stem and root ultrastructure are discussed in relation to the degree of salt resistance of these three species. Our results suggest that both N. retusa and A. halimus show high tolerance to salinity while M. arborea was considered as a salt tolerant species.     

Comparative petiole anatomy of the tribe Sorbarieae (Rosaceae) provide new taxonomically informative characters

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.     

Transformation of pollen embryo-derived explants by <Emphasis Type="Italic">Agrobacterium tumefaciens</Emphasis> in <Emphasis Type="Italic">Hyoscyamus niger</Emphasis>

Leaf, root, stem, petiole, hypocotyl, and zygotic embryo explants, as well as pollen embryoids, and redifferentiated tissues from pollen embryoid-derived plantlets of Hyoscyamus niger L. (black henbane) were inoculated with Agrobacterium tumefaciens, harboring binary vectors (pGS Gluc1) and then cultured on media containing kanamycin. Transient -glucuronidase activity and kanamycin resistant callus formation were influenced by explant origin. Transgenic calluses were obtained at a frequency of up to 30% from all the explants tested. However, transgenic shoots were obtained only from the hypocotyl of plantlets derived from pollen embryoids. Transformation was confirmed by the ability of leaf segments to produce kanamycin resistant calluses, -glucuronidase histochemical and flurometric assays, polymerase chain reaction and Southern blot analysis. The results show that pollen embryoid-derived explants may be an alternative source for both efficient transformation and regeneration of transgenic plants in recalcitrant species.     

COMPARATIVE STUDIES OF THE NODAL AND VASCULAR ANATOMY IN THE NEOTROPICAL CYATHEACEAE. I. METAXYA AND LOPHOSORIA

Comparative studies of the nodal and vascular anatomy in the monotypic genera Metaxya and Lophosoria are discussed as they relate to the taxonomy and phylogeny of the Cyatheaceae. Both genera are distinctive and primitive with respect to habit, stem and petiole indument, stelar pattern, and nodal anatomy. Metaxya possesses a prostrate, dorsiventral rhizome, whereas a short, upright radial stem occurs in Lophosoria. Trichomes occur on the stems and leaf petioles of these genera. Both Metaxya and Lophosoria have a spiral phyllotaxy, and adventitious buds occur on the petiole bases. The stelar pattern is basically a siphonostele, although frequently a dictyostele is found in Lophosoria. Accessory bundles are lacking in both genera. A characteristic petiole pattern is found in these genera, with an increase in complexity from an undivided strand in Metaxya to the three-parted petiole pattern in Lophosoria. Data from nodal and vascular anatomy indicate that these taxa are distinct from the other genera in the Cyatheaceae and belong in an independent position at the base of the Cyatheoid line, although in some respects an affinity to members of the Dicksoniaceae is indicated.     

VASCULAR DYSFUNCTION IN FUSARIUM WILT OF TOMATO

Leaves on tomato plants infected with Fusarium oxysporum f. lycopersici frequently wilt unilaterally when the vascular bundles supplying the affected leaflets are diseased. However, when the vascular bundles on one side of healthy petioles are severed by notching the petiole base, the entire leaf remains turgid. Leaflets on the notched side receive water by diffusion between bundles at the petiole tip. Lateral translocation of water out of individual vessels and between bundles in diseased xylem is impaired by the impregnation of vessel walls, intercellular spaces, and cells adjacent to vessels with the products of vascular discoloration. Waterproofing of vascular bundles can be induced in excised healthy leaves by culture filtrates of the pathogen and catechol. Waterproofing of vessels may play an important role in vascular dysfunction by confining water to individual vessels and thereby increasing the importance of vessel occlusions.     

Indole-3-acetic acid accumulation during poplar rhizogenesis revealed by immunohistochemistry

Poplar hybrid 741 [Populus alba × (P. davidiana + P. simonii) × P. tomentosa] leaves were rooted within 8 d when cultured in vitro on 1/2 Murashige and Skoog (MS) medium. The spatial distribution of endogenous indole-3-acetic acid (IAA) in the rhizogenesis was investigated, using an immunohistochemical approach. In addition, the effect of 2,3,5-triiodobenzoic acid (TIBA) on IAA distribution was also analyzed. The results showed that a strong IAA signal was detected in the vascular bundles of the basal regions of the petioles 3 d after root induction. Furthermore, the signal in vascular bundles of the basal regions of the petioles was stronger than that of the middle regions of the petioles. Application of TIBA on lamina delayed both the accumulation of IAA in the vascular bundles and rhizogenesis. These data indicate that an endogenous IAA rise in vascular bundles is among the first signals leading to the rhizogenesis, and that it results from transportation of the hormone from the lamina of the leaf to the base of the petiole, rather than by in situ IAA generation.     

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.