Structure and ontogeny of successive cambia in Tetrastigma (Vitaceae), the host plants of Rafflesiaceae

Successive cambia in Vitaceae have been reported solely for Tetrastigma, a diverse genus of lianas found primarily in tropical Asia, extending into Australia. However, the structure and origin of these successive cambia have never been fully studied. Here we report the presence of this cambial variant in Tetrastigma retinervum and T. voinierianum, and describe its ontogeny in detail in the latter. New cambia appear successively in stems of Tetrastigma differentiating from the innermost parenchyma cells of the primary phloem, which are located interior to the pericyclic fiber strands. This study constitutes the first report of successive cambia being derived from primary phloem parenchyma in woody plants as a whole. Both species are members of Tetrastigma clade VI, the most species‐rich lineage within the genus. The examination of mature stems of additional species of Tetrastigma should determine the distribution of this unique type of cambial variant in the genus and enhance our understanding of the adaptive significance of this unusual character.     

ANOMALOUS GROWTH AND VEGETATIVE ANATOMY OF SIMMONDSIA CHINENSIS

The anatomy of the stem, root, and leaf of Simmondsia chinensis (Link) Schneider was investigated, as well as the mode of tissue formation in the stem. Perivascular tissue is present as part of the primary body; outermost cell layers of this tissue mature as a fibrous sheath. The first short-lived extrafascicular cambium is generated within the remaining parenchymatous perivascular tissue. Successive independent extrafascicular cambia, organized as complete rings or large arcs, arise within peripheral conjunctive parenchyma produced by previous cambia. Extrafascicular cambia produce secondary xylem centripetally and conjunctive tissue bands and strands of secondary phloem centrifugally. Conjunctive tissue initials produce raylike structures of conjunctive tissue; true vascular rays are absent. The phellogen is actually a region of transition where the peripheral conjunctive parenchyma of previous extrafascicular cambia undergoes further cellular subdivision; a true phellogen is lacking. Xylem bands do not represent annual or seasonal growth increments, and secondary growth in Simmondsia is an unequivocal example of the “concentric” anomaly.     

Successive cambia in Aizoaceae: products and process

The transverse and longitudinal sections of the stems and roots of 11 genera of Aizoaceae, representing a wide range of growth forms from hard fibrous stems to fibre‐free roots, were studied using light microscopy and scanning electron microscopy. In most of the genera, fibres are the first xylary product of each vascular cambium, followed by vessels in a parenchyma background. Variations on this pattern help to prove that fibres are produced by vascular cambia, except in Ruschia and Stayneria, in which both the lateral meristem and the vascular cambia produce fibres. Cylinders of conjunctive tissue parenchyma that alternate with the vascular cylinders are produced by the lateral meristem. The concept that the lateral meristem gives rise to the vascular cambia and secondary cortex is supported by photographic evidence. Radial divisions occur in the origin of the lateral meristem, and then again as vascular cambia arise from the lateral meristem; these radial divisions account for storeying in fibres and conjunctive tissue. Raylessness characterizes all Aizoaceae studied, with the exception of Tetragonia, which also differs from the remaining genera by having vasicentric axial parenchyma, a scattering of vessels amongst fibres, and the presence of druses instead of raphides. Several vascular cambia are typically formed per year. Several vascular cambia are active simultaneously in a given stem or root. Roots have fewer fibres and more abundant conjunctive tissue parenchyma than stems. Successive cambia result in an ideal dispersion of vascular tissue with respect to water and photosynthate storage and retrieval capabilities of the parenchyma, and to liana stem plans. The distribution and relative abundance of fibres, vessels, secondary phloem, and conjunctive tissue parenchyma relate primarily to habit and are not a good source of systematic data, with the probable exception of Tetragonia. The general pattern of lateral meristem and vascular cambial ontogeny is the same as in other families of the core Caryophyllales, although the patterns of the tissues produced are diverse. © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 153 , 141–155.     

The development of phloem anastomoses between vascular bundles and their role in xylem regeneration after wounding in Cucurbita and Dahlia

The differentiation of phloem anastomoses linking the longitudinal vascular bundles has been studied in stem internodes of Cucurbita maxima Duchesne, C. pepo L. and Dahlia pinnata Cav. These anastomoses comprise naturally occurring regenerative sieve tubes which redifferentiate from interfascicular parenchyma cells in the young internodes. In all three species, severing a vascular bundle in a young internode resulted in regeneration of xylem to form a curved by-pass immediately around the wound. The numerous phloem anastomoses in these young internodes were not involved in this process, the regenerated vessels originating from interfascicular parenchyma alone. Conversely, in mature internodes of Dahlia, the regenerated vessels originated from initials of the interfascicular cambia, and their phloem anastomoses did not influence the pattern of xylogenesis. On the other hand, in old internodes of Cucurbita, in which an interfascicular cambium was not yet developed, the parenchyma cells between the bundles had lost the ability to redifferentiate into vessel elements, and instead, regenerated vessels were produced in the phloem anastomoses. Thus, the wounded region of the vascular bundle was not bypassed via the shortest, curved pathway, but by more circuitous routes further away from the wound. Some of the regenerated vessels produced in the phloem anastomoses were extremely wide, and presumably efficient conductors of water. It is proposed that the dense network of phloem anastomoses developed during evolution as a mechanism of adaptation to possible damage in mature internodes by providing flexible alternative pathways for efficient xylem regeneration in plants with limited or no interfascicular cambium.This paper is dedicated to the memory of the late Isaac Blachmann (deceased 19 November 1995), father-in-law of the senior author, for encouragement and advice throughout the yearsThis research was supported by an International Scientific Exchange Award to R.A. from the Israel Academy of Sciences and The Royal Society.     

ANATOMY OF NODES VS. INTERNODES IN COLEUS: THE NODAL CAMBIUM

Secondary growth begins in the nodal regions before the internodal regions in Coleus, so that longitudinally discontinuous vascular cambia are formed in the 6th through the 9th or 10th nodes, where the internodal cambium becomes continuous between nodal cambia. The nodal cambia are identifiable by radial seriation in interfascicular regions, typical cytology of fusiform initials, and the presence of a ray system. Anatomical features distinct from the primary plant body are shared by the nodal and internodal cambia. Branching of primary vascular strands, restricted to procambium and phloem, is virtually confined to nodal regions. In secondary growth, vascular branching of xylem and phloem occurs in both nodes and internodes. Xylem strand branches are formed only from derivatives of vascular cambia. It is proposed that the cambium provides the secondary plant body an efficient channel for lateral auxin transport, by which branching across interfascicular regions is facilitated.     

Comparative wood anatomy ofCoriaria

Wood anatomy ofCoriaria was surveyed to clarify generic features on the basis of 14 species collected from various regions of the World to cover the whole range of geographic distribution and habitual variation. Wood anatomy ofcoriaria is considerably uniform, and the species share a combination of the following features: 1) pores are thin-walled, polygonal in outline and mostly in multiples; 2) vessel elements and libriform fibers are very short; 3) perforation plates are exclusively simple; 4) intervessel pits are alternate; 5) vascular tracheids are present; 6) wood parenchyma is vasicentric and sometimes confluent; 7) rays are heterogeneous and large. Its species differ in several characters, such as distinctness of growth rings, pore size, pore patterns, type and abundance of wood parenchyma, and distinctness of storied structure. Comparisons among species indicate that the species of the Northern Hemisphere show a tendency toward having semi-ring porosity, while those of the Southern and Western Hemisphere have diffuse porosity. The other infrageneric variations appear to be related to different habits of the species rather than to geographic distribution. Small trees mostly have confluent and vasicentric parenchyma composed of fusiform cells and distinctly storied tissues, while shrubs and herbs have less abundant parenchyma which is vasicentric and comprises strands of two to four cells and indistinctly storied tissues.     

Radial secondary growth and formation of successive cambia and their products in Ipomoea hederifolia L. (Convolvulaceae)

Ipomoea hederifolia stems increase in thickness using a combination of different types of cambial variant, such as the discontinuous concentric rings of cambia, the development of included phloem, the reverse orientation of discontinuous cambial segments, the internal phloem, the formation of secondary xylem and phloem from the internal cambium, and differentiation of cork in the pith. After primary growth, the first ring of cambium arises between the external primary phloem and primary xylem, producing secondary phloem centrifugally and secondary xylem centripetally. The stem becomes lobed, flat, undulating, or irregular in shape as a result of the formation of both discontinuous and continuous concentric rings of cambia. As the formation of secondary xylem is greater in one region than in another, this results in the formation of a grooved stem. Successive cambia formed after the first ring are of two distinct functional types: (1) functionally normal successive cambia that divide to form secondary xylem centripetally and secondary phloem centrifugally, like other dicotyledons that show successive rings, and (2) abnormal cambia with reverse orientation. The former type of successive rings originates from the parenchyma cells located outside the phloem produced by previous cambium. The latter type of cambium develops from the conjunctive tissue located at the base of the secondary xylem formed by functionally normal cambia. This cambium is functionally inverted, producing secondary xylem centrifugally and secondary phloem centripetally. In later secondary growth, xylem parenchyma situated deep inside the secondary xylem undergoes de‐differentiation, and re‐differentiates into included phloem islands in secondary xylem. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 30–40.     

The Developmental Anatomy of the Root System in Yam Bean, Pachyrhizus erosus Urban

Investigations revealed that the anatomy of the primary radicularroot of yam bean (Pachyrhizus erosus L.) was typically dicotyledonousexcept that the xylem was not completely developed centripetally.Most of the roots had tetrarch xylem, although a few triarchand pentarch roots were also observed. In both tuberous andnon-tuberous roots, secondary thickening occurred by the formationof the meristematic vascular cambium which formed secondarytissues in a normal fashion. Subsequently, tuberization wasinitiated in the secondary xylem by the development of anomalous‘secondary’ cambia from parenchyma cells surroundingvessel elements. Anomalous ‘secondary’ cambia alsodeveloped from parenchyma cells not associated with vessels.Subsequently, anomalous ‘tertiary’ cambia differentiatedfrom tissues produced by the anomalous ‘secondary’cambia. Activities of these anomalous cambia resulted in theproduction of parenchyma storage cells and were chiefly responsiblefor the growth of the mature tuber. Pachyrhizus erosus L., yam bean, tuberous root, anatomy, anomalous ‘secondary’ cambia, anomalous ‘tertiary’ cambia, centripetal xylem development     

Continuity of Procambium and Anomalous Cambium During Formation of Successive Cambia in Celosia argentea

The development of woody plants is related to the continuity of the procambium and cambium. Whether such a continuity is present in plants with successive cambia, especially in those, where the first cambium is formed outside the primary vascular bundles, has not been analyzed so far. Therefore, we studied the development of vascular meristem in Celosia argentea, in which the first and successive cambial cylinders arise outside the primary bundles and, intriguingly, in the literature are interpreted as developmentally independent structures. Our results showed that in C. argentea, the outermost procambial cells maintain their meristematic characteristics during differentiation of vascular bundles and divide periclinally, forming the zone of procambium-derived cells outside the primary bundles. This zone comprises parenchyma cells bordering the bundles, and a continuous ring of the incipient cambial cells neighboring the primary cortex. Later in the development, the ability to preserve the outermost cells in the cambium undifferentiated is repeated during the formation of successive cylinders of cambia. Together, our results clearly point to the developmental continuity of the procambium and successive cambia in C. argentea, despite their seemingly spatial distinctiveness. We postulate that the mechanism demonstrated in C. argentea is universal and orchestrates the development of successive cambia in other plant species.

     

Secondary root growth in Rhynchosia edulis Griseb. (Leguminosae): Origin of cambia and their products

This study focuses on the development of a secondary root structure in Rhynchosia edulis Griseb. (Leguminosae). Its principal objectives are (i) to study the origin of cambia and the nature of their products; (ii) to correlate root structure to habitat; and (iii) to compare this anatomy with that of other Leguminosae species growing in the same environment. Serial transverse cuts of the main root show that the secondary root structure in this species results from several phenomena, namely (1) a cambium arising from procambial and pericycle cells; (2) a lateral meristem producing cell layers from the periphery towards the inner part of the root and from which vascular bundles, whose cambia fuse forming a continuous ring, originate; and (3) the formation of “elliptical cambia” in the mid portion of the root giving rise to vascular bundles in reverse orientation. The comparison of secondary root growth in R. edulis with other root structures in Leguminosae species growing in hilly areas shows different structural patterns. Nonetheless, these different patterns have the same objective: to enlarge storage parenchyma tissue enabling survival within an environment having limited water availability.     

SUCCESSIVE CAMBIA IN THE STEM OF PHYTOLACCA DIOICA

Phytolacca dioica L., an evergreen tree of the Phytolaccaceae, is one of the species of Phytolacca which shows anomalous secondary thickening in its stem. This mode of thickening has been regarded as successive cambial activity or alternatively, in some more recent interpretations, as thickening by unidirectional activity of a cambial zone. The stem thickening of P. dioica is of the former type. The cambium produces fascicular strands, showing centrifugal differentiation of xylem and centripetal differentiation of phloem on opposite sides of the cambial layer, and rays are produced between the fascicular areas. In both xylem and phloem the younger elements are closer to the cambium than the older elements. Succeeding cambia arise periodically by periclinal divisions in a layer of parenchyma cells two or three cells beyond the outermost intact phloem derived from the current cambium. Each cambium forms a few parenchyma cells on both sides before it forms derivatives which mature into lignified xylem elements or conductive elements of the phloem. The parenchyma thus formed toward the outside later becomes the site of the origin of the succeeding cambium. Only one or two layers of this phloem parenchyma go on to form the new cambium; the remaining cells accumulate between the outermost phloem and the cortex. P. weberbaueri shows stem structure similar to P. dioica. P. meziana, a shrub, shows normal stem structure.     

Nodulation of white clover (Trifolium repens) in the absence ofRhizobium

Summary Spontaneous nodules developed on the roots of white clover (Trifolium repens cv. Ladino) in the absence ofRhizobium. A small subpopulation of uninoculated clover plants (0.2%) exhibited white, single-to-multilobed elongated structures on their root systems when grown without fixed nitrogen. Clonal propagation using aseptic stolons confirmed the genetic stability of the observation. Few if any viable bacteria of unknown origin were recovered from surfacesterilized structures. Nodule contents were incapable of eliciting nodulation. Histological observations showed that these structures possessed all the characteristic features of indeterminate nodules, such as active meristem, cortex, endodermal layer, vascular strands, and a central zone with parenchyma cells. Infection threads, intercellular or intracellular bacteria were absent. Instead, numerous starch grains were observed in the central zone, a feature absent in normal nitrogen-fixing nodules. Our observation broadens the concept of spontaneous nodulation, believed to be restricted to alfalfa (Medicago sativa), to other legumes, and suggests a degree of generality among indeterminately nodulated legumes displaying natural heterozygosity.     

Morphogenesis of <Emphasis Type="Italic">Paragordius varius</Emphasis> (Nematomorpha) during the parasitic phase

The development of Paragordius varius (Nematomorpha), in the parasitic phase of the life cycle, was followed ultrastructurally from 10 days after controlled infection of crickets (Gryllus firmus) to mature animals at 30 days after infection. During this time span, specimens grow from about 1 cm to more than 10 cm and from 93 m to about 400 m in diameter. A thin larval cuticle is replaced at about day 20 by a robust adult cuticle. Epidermis and intestine appear physiologically active during the early stages, but decrease in size and cytological components during further development. This is probably connected to the uptake of nutrients through the larval cuticle, whereas the adult cuticle has only protective function. The longitudinal musculature grows continuously and changes from fewer platymyarian cells to abundant coelomyarian cells. The ventral nerve cord shifts from an intraepidermal to a submuscular position during development. Additionally, basiepidermal nerves are present. Gonads develop from paired dorsolateral compact strands. It seems that in males, cells within these strands separate into epithelial and central cells, which become the gametes, while in females, gametes proliferate from the strands into the primary body cavity and fill it up completely. In males, there is a large quantity of parenchyma, restricting the body cavity to a small periintestinal space, while in females, parenchyma occurs only in the periphery of the gametes.     

Interxylary phloem: Diversity and functions

Interxylary phloem is here defined as strands or bands of phloem embedded within the secondary xylem of a stem or root of a plant that has a single vascular cambium. In this definition, interxylary phloem differs from intraxylary phloem, bicollateral bundles, pith bundles, and successive cambia. The inclusive but variously applied terms included phloem and internal phloem must be rejected. Histological aspects of interxylary phloem are reviewed and original data are presented. Topics covered include duration of interxylary phloem; relationship in abundance between sieve tubes in external phloem and interxylary phloem; distinctions between interxylary and intraxylary phloem; presence of parenchyma, fibers, and crystals in the interxylary phloem strands; development of cambia within interxylary phloem strands; three-dimensionalization and longevity of phloem, systematic distribution of interxylary phloem; physiological significance; and habital correlations. No single physiological phenomenon seems to explain all instances of interxylary phloem occurrence, but rapidity and volume of photosynthate transport seem implicated in most instances.     

ORIGIN AND ACTIVITY OF SUCCESSIVE CAMBIA IN CYCAS (CYCADALES)

Observations of the vascular tissue of Cycas shoots have provided supporting evidence that the first vascular cambium as well as subsequent successive cambia are simultaneously active. The establishment of the second cambium occurs during the seedling stage, and differentiates mainly within the cortical cells. However, cambial activity also occurs within phloem parenchyma cells of the first vascular cylinder. Tracheids in the first and the successive vascular cylinders are generally of the same length; however, there is a trend toward increasing length within the successive cylinders, possibly because the successive cambia are long-lived.     

Development of successive cambia and formation of flat stems in Rhynchosia pyramidalis (Lam.) Urb. (Fabaceae)

Stem flattening in Rhynchosia pyramidalis (Fabaceae) is achieved by the development of crescent-shaped successive cambia on two opposite sides of the stem (referred hereafter as distal side). Other lateral sides of the stem (adjacent to supporting host and its opposite side, referred as proximal sides) usually possess single cambium. In the young stems, parenchymatous cells located outside to protophloem of distal side dedifferentiate and develop small segments of cambium. Concomitant to bidirectional differentiation of the secondary xylem and phloem, these newly developed cambial segments also extend in tangential directions. Differential activity of newly developed crescent-shaped cambial segments deposits more secondary xylem at median position as compared to their terminal ends of the stem on distal side; consequently, it pushes the cambial segment outside, thus resulting in crescent-shaped arcs of the cambia only on two opposite sides. After the production of 1–2 mm of secondary xylem, they cease to divide and new segments of cambial arc develop on the same side in a similar fashion. Such repeated behaviour of successive cambia development consequently leads to the formation of tangentially flat stems. The secondary xylem is diffusely porous with indistinct growth rings and is composed of vessels (wide and narrow), fibres, axial ray parenchyma cells, while phloem consisted of sieve elements, companion cells, axial and ray parenchyma. Rays in both xylem and phloem are uni- to multiseriate and heterocellular. The structure of secondary xylem and development of successive cambia is correlated with climbing habit.     

药用植物川牛膝根中异常次生结构的发育解剖学研究

药用植物川牛膝的根内具有异常的次生结构。其异常的次生生长是由维管柱外围发生的异常形成层通过正常的活动方式完成的。后一轮异常形成层起源于前一轮异常形成层向外产生的薄壁组织细胞,位于韧皮部的外侧。每一轮异常形成层向内产生木质部,向外产生韧皮部,组成异常维管束。其中,木质部最先开始分化。异常维管束排成螺旋状,分散在结合组织中。除最外轮一些木质部束之间的结合组织是厚壁组织外,其余结合组织都是薄壁的。由于初生结构和早期的次生结构是正常的,所以,这种异常结构可能是后起的特征。     

Ecological significance of wood anatomy in two lianas from arid southwestern Saudi Arabia

The hot and arid lowlands of southwestern Saudi Arabia are home to two common lianas, Cocculus pendulus and Leptadenia arborea. This paper attempts to relate the adaptation of these two climbing woody perennials to such a harsh environment to the anatomy and hydraulic characteristics of their wood. The stems of these lianas have wood with wide xylem vessels and high hydraulic conductivity which should enhance water flow to the upper canopy despite their severe twisting. Hydraulic conductivity is further helped by the simple perforation plates of xylem vessels. The circular thickening of xylem walls gives them strength and reduces the risk of their collapse and the ensuing embolism in the advent of high tension created by severe water deficit and high evapo-transpiration demand. Wide vessels, on the other hand, are more susceptible to embolism. This problem may be overcome by reducing the solute potential of xylem sap by hydrolysis of starch grains which were found to be abundant in the vicinity of the vessels. This should help absorb water by the deep roots from the capillary fringes of the typically shallow water table in this particular habitat. Furthermore, the abundance of ray parenchyma cells between xylem groups of both lianas provides great flexibility with minimum damage to water conduits in the stem during climbing and twisted growth. It was concluded that these wood features in both lianas are crucial for survival under the harsh conditions of arid Tihama plains of southwestern Saudi Arabia.     

Contributions to the Wood Anatomy of the Rubioideae (Rubiaceae)

Damnacanthus , Lasianthus, Saldinia, and Trichostachys are also included. Wood anatomical characters are compared with recent phylogenetic insights into the study group on the basis of molecular data. The observations demonstrate that the delimitation and separation of several taxa from the former Coussareeae/Morindeae/Prismatomerideae/Psychotrieae aggregate is supported by wood anatomical data. The Coussareeae can be distinguished from the other Rubioideae by their scanty parenchyma, septate libriform fibres, and the combination of uniseriate and very high multiseriate rays with sheath cells. Axial parenchyma bands and fibre-tracheids characterise Gynochtodes and some species of Morinda (Morindeae s.str.), but the latter genus is variable with respect to several features (e.g. vessel groupings and axial parenchyma distribution). Wood data support separation of Rennellia and Prismatomeris from Morindeae s.str.; vessels in both genera are exclusively solitary and axial parenchyma is always diffuse to diffuse-in-aggregates. Damnacanthus differs from the Morindeae alliance by the occurrence of septate fibres, absence of axial parenchyma, and the occasional presence of fibre wall thickenings. There are interesting similarities between members of the Lasianthus clade and the Pauridiantheae/Urophyleae group such as the sporadic occurrence of spiral thickenings in axial parenchyma cells. Received 26 January 2001/ Accepted in revised form 6 June 2001