Seasonal development of secondary xylem and phloem in <Emphasis Type="Italic">Schizolobium parahyba</Emphasis> (Vell.) Blake (Leguminosae: Caesalpinioideae)

The cambial activity and periodicity of secondary xylem and phloem formation have been less studied in tropical tree species than in temperate ones. This paper describes the relationship between seasonal cambial activity, xylem and phloem development, and phenology in Schizolobium parahyba, a fast growing semideciduous seasonal forest tree from southeastern Brazil. From 2002 to 2003, wood samples were collected periodically and phenology and climate were recorded monthly in the same period. S. parahyba forms annual growth increments in wood, delimited by narrow initial parenchyma bands. The reduction of the cambial activity to a minimum correlates to the dry season and leaf fall. The higher cambial activity correlates to the wet season and the presence of mature leaves. In phloem, a larger conductive region was observed in the wet season, when the trees were in full foliage. The secondary phloem did not exhibit any incremental zone marker; however, we found that the axial parenchyma tends to form irregular bands.     

Gibberellin signaling

A study of stem anatomy and the sclerenchyma fibre cells associated with the phloem tissues of hemp (Cannabis sativa L.) plants is of interest for both understanding the formation of secondary cell walls and for the enhancement of fibre utility as industrial fibres and textiles. Using a range of molecular probes for cell wall polysaccharides we have surveyed the presence of cell wall components in stems of hemp in conjunction with an anatomical survey of stem and phloem fibre development. The only polysaccharide detected to occur abundantly throughout the secondary cell walls of phloem fibres was cellulose. Pectic homogalacturonan epitopes were detected in the primary cell walls/intercellular matrices between the phloem fibres although these epitopes were present at a lower level than in the surrounding parenchyma cell walls. Arabinogalactan-protein glycan epitopes displayed a diversity of occurrence in relation to fibre development and the JIM14 epitope was specific to fibre cells, binding to the inner surface of secondary cell walls, throughout development. Xylan epitopes were found to be present in the fibre cells (and xylem secondary cell walls) and absent from adjacent parenchyma cell walls. Analysis of xylan occurrence in the phloem fibre cells of hemp and flax indicated that xylan epitopes were restricted to the primary cell walls of fibre cells and were not present in the secondary cell walls of these cells.     

Structure and ultrastructure of leaf and calyx glands in Galphimia brasiliensis (Malpighiaceae)

The present study describes the anatomical structure of calyx and leaf glands in Galphimia brasiliensis and analyzes the mechanism of secretion. The glands are marginal and suprabasal, cup-shaped, sessile, and scarcely visible with the naked eye. Light microscopy reveals the following features: a thin, smooth cuticle; unistratified secretory cells; subglandular parenchyma; and vascular bundle supply composed of phloem and xylem with abundant druses of calcium oxalate. Transmission electron microscopy reveals the presence of secretory cells with conspicuous nuclei, dense cytoplasm, lipid droplets, numerous vesicles, mitochondria, Golgi, rough endoplasmic reticulum (RER), and elongated plastids with osmiophilic contents. The secretion reaches the apoplastic space and accumulates beneath the cuticle. Finally, the viscous, translucent exudate is eliminated by mechanical rupture of the cuticle. Histochemical analysis confirms that lipids are the main constituent. Small amounts of polysaccharides were also identified.     

Stem anatomy of Dolichos lablab Linn (Fabaceae): Origin of cambium and reverse orientation of vascular bundles

Secondary growth in the stem of Dolichos lablab is achieved by the formation of eccentric successive rings of vascular bundles. The stem is composed of parenchymatous ground tissue and xylem and phloem confined to portions of small cambial segments. However, development of new cambial segments can be observed from the obliterating ray parenchyma, the outermost phloem parenchyma and the secondary cortical parenchyma. Initially cambium develops as small segments, which latter become joined to form a complete cylinder of vascular cambium. Each cambial ring is functionally divided into two distinct regions. The one segment of cambium produces thick-walled lignified xylem derivatives in centripetal direction and phloem elements centrifugally. The other segment produces only thin-walled parenchyma on both xylem and phloem side. In mature stems, some of the axial parenchyma embedded deep inside the xylem acquires meristematic activity and leads to the formation of thick-walled xylem derivatives centrifugally and phloem elements centripetally. The secondary xylem comprises vessel elements, tracheids, fibres and axial parenchyma. Rays are uni-multiseriate in the region of cambium that produces xylem and phloem derivatives, while in some of the regions of cambium large multiseriate, compound, aggregate and polycentric rays can be noticed.     

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.     

Anatomy of the floral nectaries of some neotropical Salacioideae (Celastraceae)

Floral nectaries have contributed to the systematics of different taxonomic groups. Since those of the neotropical genera included in subfamily Salacioideae—Cheiloclinium Miers, Peritassa Miers, Salacia L. and Tontelea Aubl.—have different forms and positions, we explored their anatomy to delimit more precisely the genera of subfamily Salacioideae. Buds and open flowers of six species were treated following the usual techniques in plant anatomy. The obtained data were helpful in characterizing the floral nectary anatomy of the studied species. Furthermore, some features such as form, position and surface of nectaries; form of their epidermal cells; presence and distribution of stomata; occurrence of idioblasts containing druses in the nectariferous parenchyma; and absence of nectary vascularization can contribute to the taxonomy and phylogeny of the Salacioideae studied. In most of the studied species the nectar is probably released by both the stomata and the nectary epidermal surface. In Cheiloclinium cognatum, the structure acknowledged as nectary is actually a vestigial tissue and the functions of attracting and rewarding pollinators has phylogenetically migrated to the stigmatic region. The druses and phenolic substances observed in the nectariferous parenchyma probably help defend flowers against herbivore attacks. The minute size of the nectaries of Salacioideae may explain the absence of vascularization. The floral nectaries of Salacia elliptica are epithelial while those of the other species are mesenchymal.     

Comparative anatomy and systematics of Catasetinae (Orchidaceae)

Catasetinae consist of five genera of pseudobulbous Orchidaceae of the Neotropics. Anatomy is characterized by sunken, three-celled foliar hairs, mostly tetracytic stomatal apparatuses, superficial stomata, homogeneous mesophyll, foliar fibre bundles, collateral vascular bundles in a single row, xylem and phloem sclerenchyma associated with vascular bundles in leaves, conical, and rough-surfaced silica bodies adjacent to vascular bundle sclerenchyma; epidermal cells of pseudobulbs with heavily thickened outer walls, pseudobulb ground tissue of assimilatory and water-storage cells, scattered vascular bundles in pseudobulbs, and sclerenchyma and stegmata associated only with phloem of pseudobulbs; roots with thin-walled velamen cells and tenuous spirals of cell wall material, distinctive epivelamen cells, thin-walled exodermal cells and vascular tissue embedded in parenchyma. Except for mucilaginous idioblasts that occur in Mormodes and Cycnoches , there are few outstanding anatomical differences among the five genera. Thus, there are few anatomical characteristics of phylogenetic value. The monophyly of Catasetinae is supported by the presence of sunken foliar hairs. Our results support a close relationship between Clowesia and Catasetum , and between Mormodes and Cycnoches. Among the outgroups Pteroglossaspis is especially distinctive.     

EXPERIMENTAL INDUCTION OF VASCULAR TISSUES IN CALLUS OF ANGIOSPERMS

Wetmore , Ralph H. (Harvard U., Cambridge, Mass.), and John P. Rier . Experimental induction of vascular tissues in callus of angiosperms . Amer. Jour. Bot. 50(5): 418–430. Illus. 1963.—Callus tissues in established maintenance culture lack morphological and physiological organization. Such callus consists of homogeneous parenchyma. Movement of auxin and sugar, therefore, must be along diffusion gradients. The only vascular tissues occurring in callus are induced. Experimental induction of vascular tissues has been successful in callus of 3 sp. of the Oleaceae: a tree, Fraxinus americana, and 2 shrubs, Syringa vulgaris and Ligustrum vulgare; another tree, Salix purpurea, var. lambertiana; a vine, Parthenocissus tricuspidata; and an herb, Helianthus tuberosus. In each of these species, an auxin (IAA or NAA in these studies) and a sugar (sucrose or glucose in these studies) prove necessary for the induction and complete differentiation of xylem and phloem in callus tissues. Varying concentrations of sugar alter the proportions of xylem to phloem: low concentrations, 1.5%–2.5%, favor xylem formation; high, 3%–4%, favor phloem. Middle concentrations, 2.5%–3.5%, favor the presence of xylem and phloem, usually with a cambium between. The almost universal association of xylem and phloem may have its explanation in this middle concentration of sugar. Grafting of apices into callus or direct application of appropriate concentrations of an auxin and a sugar in agar to the surface of callus causes nodules of vascular tissue to be formed, mostly in a circular pattern when seen in section transverse to the axis of orientation of the callus in the medium. The diameter of this circle varies directly with the auxin concentration at the place of application, 0.05 mg/liter giving a narrow, and 1 mg/liter, a wide pith. In individual nodules, xylem is characteristically oriented towards the center of the callus and the phloem towards the outside. Variable cross-sectional views of nodule distribution in calli under different treatments suggest experimental approaches to understanding stelar patterns. The induction and differentiation in callus of xylem and phloem tissues has no relation to conduction. Any use of vascular tissues can occur only after their induction.     

Stem anatomy of the dwarf subshrub Cressa cretica L. (Convolvulaceae)

Stem anatomy and development of medullary phloem are studied in the dwarf subshrub Cressa cretica L. (Convolvulaceae). The family Convolvulaceae is dominated by vines or woody climbers, which are characterized by the presence of successive cambia, medullary- and included phloem, internal cambium and presence of fibriform vessels. The main stems of the not winding C. cretica shows presence of medullary (internal) phloem, internal cambium and fibriform vessels, whereas successive cambia and included phloem are lacking. However, presence of fibriform vessels is an unique feature which so far has been reported only in climbing members of the family. Medullary phloem develops from peri-medullary cells after the initiation of secondary growth and completely occupies the pith region in fully grown mature plants. In young stems, the cortex is wide and formed of radial files of tightly packed small and large cells without intercellular air spaces. In thick stems, cortical cells become compressed due to the pressure developed by the radial expansion of secondary xylem, a feature actually common to halophytes. The stem diameter increases by the activity of a single ring of vascular cambium. The secondary xylem is composed of vessels (both wide and fibriform), fibres, axial parenchyma cells and uni-seriate rays. The secondary phloem consists of sieve elements, companion cells, axial and ray parenchyma cells. In consequence, Cressa shares anatomical characteristics of both climbing and non-climbing members. The structure of the secondary xylem is correlated with the habit and comparable with that of other climbing members of Convolvulaceae.     

Cambial Activity and Distribution of Rubber in Guayule (Parthenium argentatum)

Vascular cambium in Guayule, a rubber producing Mexican shrubof Asteraceae family is non-storied. Cambial activity variesperiodically, and the vascular cambium and its immediate derivativesdo not contain rubber. However, as the xylem and phloem parenchymacells derived from the vascular cambium age, rubber depositionstarts from the cell periphery along the walls and later towardstheir cell lumen. Though the sieve tubes and companion cellsof phloem contain no rubber, all parenchyma cells of xylem andphloem, show the presence of rubber, though its amount varies.However, certain lignified xylem ray cells and lignified pithcells are devoid of rubber accumulation. Microfluorescence studiesshow that the epithelial, phloem ray parenchyma, cortical andpith cells, in descending order, have the highest to lowestrubber content. The size and number of rubber particles observedin the parenchyma cells are greatest during the period of cambialdormancy than in an active cambial period Cambium, guayule, rubber     

PHLOEM OF SPHENOPHYLLUM

The phloem of most fossil plants, including that of Sphenophyllum, is very poorly known. Sphenophyllum was a relatively small type of fossil arthrophyte with jointed stems bearing whorls of leaves ranging in form from wedge or fan-shaped to bifid, to linear. The aerial stem systems of the plant exhibited determinate growth involving progressive reduction in the dimensions of the stem primary bodies, fewer leaves per whorl, and smaller and simpler leaves distally. The primary phloem occurs in three areas alternating in position with the arms of the triarch centrally placed primary xylem. Cells of the primary phloem, presumably sieve elements, are axially elongate with horizontal to slightly tapered end walls. In larger stems with abundant secondary xylem and secondary cortex or periderm, a zone of secondary phloem occurs whose structure varies in the three areas opposite the arms of the primary xylem, as opposed to the three areas lying opposite the concave sides of the primary xylem. The axial system of the secondary phloem consists of vertical series of sieve elements with horizontal end walls. In the areas opposite the protoxylem the parenchyma is present as a prominent ray system showing dilation peripherally. Sieve elements in the areas opposite the protoxylem arms have relatively small diameters. In the areas between the protoxylem poles the secondary phloem sieve elements have large diameters and are less obviously in radial files, while the parenchyma resembles that of the secondary xylem in these areas in that it consists of strands of cells extending both radially and tangentially. An actively meristematic vascular cambium has not been found, indicating that this layer changed histologically after the cessation of growth in the determinate aerial stem systems and was replaced by a post-meristematic parenchyma sheath made up of axially elongate parenchyma lacking cells indicative of being either fusiform or ray initials. A phellogen arose early in development in a tissue believed to represent pericycle and produced tissue comparable to phellem externally. Normally, derivatives of the phellogen underwent one division prior to the maturation of the cells. Concentric bands of cells with dark contents apparently represent secretory tissue in the periderm and cell arrangements indicate that a single persistent phellogen was present. Sphenophyllum is compared with other arthrophytes as to phloem structure and is at present the best documented example of a plant with a functionally bifacial vascular cambium in any exclusively non-seed group of vascular plants.     

太子参微块根发育的解剖学与组织化学定位

采用植物组织培养、解剖学及组织化学定位方法研究太子参试管微块根发育的形态结构与营养物质积累特征的结果表明：太子参微块根由组培苗膨大的腋芽基部长出的不定根发育而成,经历了初生结构与次生结构发育,其膨大加粗是由于不定根的次生生长。维管形成层向内形成大量的次生木质部构成微块根的主要部分。淀粉粒是太子参微块根的主要营养存储方式。随着微块根的次生生长,淀粉粒先在次生木质部薄壁细胞中形成,随后在次生韧皮薄壁细胞中也大量积累。膨大的微块根可以合成太子参皂苷,成熟微块根中次生韧皮部的皂苷含量略高于次生木质部。离体太子参微块根的生长发育和营养物质的积累与块根中的相同。     

白鲜营养器官解剖结构及其与白鲜碱的积累关系

应用植物解剖学、组织化学及植物化学方法对白鲜营养器官根、茎、叶的结构及其生物碱的积累进行了研究。结果显示:(1)白鲜根的次生结构以及茎和叶的结构类似一般双子叶植物;白鲜多年生根主要由周皮、次生韧皮部、维管形成层以及次生木质部组成,根次生韧皮部中可见大量的淀粉、草酸钙簇晶、韧皮纤维以及油细胞;茎由表皮、皮层、维管组织和髓组成;叶由表皮、栅栏组织、海绵组织和叶脉组成;在茎和叶初生韧皮部的位置均分布有韧皮纤维,在叶表皮上分布有头状腺毛和非腺毛;在茎和叶紧贴表皮处分布有分泌囊。(2)组织化学分析结果显示:在白鲜多年生根中,生物碱类物质主要分布在周皮、次生韧皮部、维管形成层和木薄壁细胞中;在茎中,生物碱主要分布在表皮、皮层、韧皮部、木薄壁细胞及髓周围薄壁细胞中;在叶中,生物碱主要分布在表皮细胞、叶肉组织和维管组织的薄壁细胞;此外在分泌囊和头状腺毛中亦含有生物碱类物质。(3)植物化学结果显示,秦岭产白鲜根皮/白鲜皮、根木质部、茎和叶中白鲜碱含量分别为0.041%、0.012%、0.004%和0.002%,其中木质部中白鲜碱含量和其他部分地区白鲜皮中白鲜碱含量类似。研究表明,在秦岭产白鲜营养器官中,除根皮/白鲜皮外,在根木质部亦含有大量的白鲜碱,且在茎和叶中亦含有一定的白鲜碱,具有潜在的开发利用价值。     

Variation in wood density and anatomy in a widespread mangrove species

Wood density is an important plant trait that influences a range of ecological processes, including resistance to damage and growth rates. Wood density is highly dependent on anatomical characteristics associated with the conductive tissue of trees (xylem and phloem) and the fibre matrix in which they occur. Here, we investigated variation in the wood density of the widespread mangrove species Avicennia marina in the Exmouth Gulf in Western Australia and in the Firth of Thames in New Zealand. We assessed how variation in xylem vessel size, fibre wall thickness and proportion of phloem within the wood contributed to variation in wood density and how these characteristics were linked to growth rates. We found the wood density of A. marina to be higher in Western Australia than in New Zealand and to be higher in taller seaward fringing trees than in scrub trees growing high in the intertidal. At the cellular level, high wood density was associated with large xylem vessels and thick fibre walls. Additionally, wood density increased with decreasing proportions of phloem per growth layer of wood. Tree growth rates were positively correlated with xylem vessel size and wood density. We conclude that A. marina can have large xylem vessel sizes and high growth rates while still maintaining high wood density because of the abundance and thickness of fibres in which vessels are found.     

DEVELOPMENT AND STRUCTURE OF THE VASCULAR CONNECTION BETWEEN THE PRIMARY AND SECONDARY ROOT OF GLYCINE MAX (L.) MERR.

The primary vascular connection between primary and secondary root of Glycine max (L.) Merr. was derived from stelar parenchyma and pericycle. Inner stelar parenchyma, associated with the parent metaxylem and outer stelar parenchyma adjacent to the pericycle, were resonsible for the histogenesis of the primary xylem connection. Acropetal maturation of the diarch xylem connection occurred after the lateral root emerged from the parent root. Development of tetrarchy occurred distal to the diarch xylem connection. The concentric primary phloem connection was derived from the pericycle and outer stelar parenchyma. Acropetal maturation of the primary phloem connection occurred prior to lateral root emergence from the parent root. Secondary growth quickly augmented the primary vascular connection. A substantial amount of mature secondary xylem formed prior to maturation of the secondary phloem. The structure of the primary and secondary vascular connections is described.     

远志根的发育解剖学研究

运用石蜡切片法对远志根的发育过程及1~3年生根的结构进行解剖学研究。结果显示:远志根的原分生组织由3群原始细胞组成,具有典型分生组织的细胞学特征。初生分生组织分化为根冠原、表皮原、皮层原和中柱原;初生结构由表皮、皮层和中柱组成,初生木质部为二原型。次生生长是依靠维管形成层和木栓形成层的活动完成,次生结构从外到内由周皮和次生维管组织组成;远志根次生结构特点为:次生韧皮部在次生维管组织中占主要部分,次生韧皮部中以韧皮薄壁细胞为主且其中储存有丰富的内含物,随着根龄的增加,韧皮薄壁细胞中的内含物也随之增加。3年生的主根中次生韧皮部薄壁细胞中的内含物最丰富;不同年份远志的主根随根龄的增加,周皮、次生韧皮部和次生木质部的面积都呈增加趋势,其中韧皮部和木质部的面积比值随根龄增长呈由小到大的变化,这是远志根的显著特点;根中的周皮发达,具有较厚的木栓层,次生木质部中导管和纤维发达,导管分布频率较高,并具有较大的口径。周皮和次生木质部的结构特征与远志的抗旱特性相适应。     

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.     

掌叶大黄根多糖的积累分布特征

采用组织化学方法和苯酚硫酸比色法研究了掌叶大黄（Rheum palmatum）根中大黄多糖的贮藏分布特征和含量变化规律。结果表明：大黄多糖在根内的贮藏是多位点的,在根周皮的栓内层、次生维管组织的薄壁细胞内不同程度地贮藏和积累了一定数量的大黄多糖,次生木质部的木薄壁细胞和次生韧皮部的韧皮薄壁细胞是主要贮藏和积累的部位;不同发育时期根中大黄多糖含量的变化规律为,随着植物的不断成熟,根及其各组织中大黄多糖的总含量表现为逐渐增高的趋势,但在发育的后期略有下降;韧皮薄壁细胞与木薄壁细胞相比,贮藏大黄多糖的含量相对较多,大黄多糖的贮藏积累方式为逐渐累积的方式。