Leaf architectural studies in the brassicaceae

The leaf architecture has been studied in 19 genera and 35 species of the Brassicaceae. The major venation pattern is pinnate craspedodromous with the exception ofAlyssum maritimum, Iberis amara, I. umbellata andMalcolmia maritima where it is pinnate-festooned brochidodromous. The number of secondary veins and their angle of divergence vary from species to species and even within the same species. Marginal ultimate venation is mostly looped occasionally incomplete. The areole size and shape is variable. The veinlets may be simple or once or twice dichotomously divided. The simple veinlets may be curved or hooked. The tracheids are either uni-, bi-, tri- or multiseriate and extraordinarily variable in size and shape. Isolated vein endings, isolated free vein endings, isolated tracheids and extension cells are observed. Myrosin cells have been observed inEruca vesicaria ssp.sativa andCochlearia cochlearioides.     

Leaf architecture of some acanthaceae

Leaf architectural pattern has been studied in 27 genera and 35 species of the Acanthaceae. The major venation pattern conforms to pinnate camptodromous with eucamptodromous or festooned brochidodromous secondaries, pinnate craspedodromous inAcanthus ilicifolius and acrodromous inLepidagathis trinervis. Intersecondary veins are common. The marginal ultimate venation is looped. The areoles are variable in size. The vein endings are usually simple, linear or cuved or divide once or twice dichotomously. Isolated vein endings, tracheids, isolated free vein endings and extension cells are observed. Transfusion tracheids are seen inDicliptera verticillata. Miniature vessel elements with simple perforation plates are noticed lying free in the areole or at the end of tracheids inSeriocalyx scaber andThunbergia grandiflora.Elytraria acaulis, Nelsonia canescens andStaurogynae zeylanica exhibit venation pattern shown by majority of the taxa studied, of the Acanthaceae.     

Leaf Architecture of Some Monocotyledons with Reticulate Venation

Twelve species belonging to seven monocotyledonous families:Hydrocharitaceae, Taccaceae, Dioscoreaceae, Smilacaceae, Araceae,Alismataceae and Aponogetonaceae show reticulate venation typicalof dicotyledons. A study of the leaves of these species showsthat venation patterns are usually curvipalmate-convergate,occasionally rectipalmate or collimate, and rarely pinnate lyratetype. Number, size and shape of areoles, number of primaries,number of secondaries along one side of the primaries, anglebetween 1 and 2° veins and number of vein endings per areoleare given for each species. Intesecondary veins, isolated tracheids,loops, extension cells, raphide and raphide idioblasts and terminaltracheids were observed. Marginal ultimate venation is mostlyarcuate. Major and minor veins are jacketed by parenchymatousbundle sheath cells. The lamina of Tacca leontopetaloides, Colocasiaesculenta and Scindapsus aureus show a single midrib-like centralregion similar to that of dicotyledonous leaves, and it is multistrandedin Aponogeton natans, Limnophyton obtusifolium and Ottelia alismoides.The degree of vein order is most commonly up to fourth or fifthand rarely up to sixth in Dioscorea hispida. Monocotyledons, leaf architecture, vein endings, venation, areoles     

Foliar architecture of vanilloid orchids: insights into the evolution of reticulate leaf venation in monocotyledons

Leaves of representative taxa within the vanilloid clade of Orchidaceae were cleared and their venation patterns studied. Within subtribe Pogoniinae, Isotria and Pogonia exhibit a prominent reticulate venation pattern, although only Pogonia ophioglossoides is characterized by free vein endings. Within subtribe Vanillinae, all species of Epistephium , as well as the New Caledonian endemics, Eriaxis rigida and Clematepistephium smilacifolium , have reticulate-veined leaves that are characterized by numerous free vein endings. Leaves of most species of Vanilla exhibit a parallel-veined pattern stereotypical of monocots, although branched secondaries with free endings were observed in V. africana. Most members of subtribe Galeolinae are 'saprophytes' with reduced bract-like leaves. Vascular bundles enter these leaf-homologues in a parallel manner, but quickly ram+ throughout the tissue in an irregular manner. Leaf venation is used to hypothesize patterns of generic relationships within the vanilloid clade. Molecular evidence for phylogenetic relationships among angiosperms indicates that reticulate leaf venation has arisen independently in several unrelated monocot families, including the vanilloid Orchidaceae, perhaps by a similar evolutionary scenario.     

内蒙古地区锦鸡儿属3种植物叶脉序比较研究

以内蒙古地区锦鸡儿属3种植物小叶锦鸡儿、中间锦鸡儿、柠条锦鸡儿为对象,利用光学显微镜观察了它们的叶脉序特征,并测定相应的数量指标.结果表明,3种锦鸡儿植物叶片在脉序类型、三级脉类型、高级脉类型、盲脉类型及分枝、网眼形态上没有明显的差别;在各级脉粗细、二级脉夹角、三级脉疏密度以及网眼发育等特征上均表现出随地理分布自东向西逐步递增、递减、密集和趋于完善的变化特点,且有交叉重叠现象;在一级脉的粗细、二级脉的分枝、二级间脉的类型、三级脉疏密等特征上,中间锦鸡儿和柠条锦鸡儿差异较小.研究发现,内蒙古地区锦鸡儿属3种植物在多数叶脉序形态特征和一些数量化指标上没有明显的差异,其亲缘关系较为紧密,但中间锦鸡儿和柠条锦鸡儿在种间关系上可能更为接近;3个种自东向西呈现很强的相似性和渐变性分布,表现出地理渐变的趋势.     

SIEVE TUBES IN FOLIAR VEIN ENDINGS: REVIEW AND QUANTITATIVE SURVEY OF RUDBECKIA LACINIATA (ASTERACEAE)

Sieve tube distribution in foliar vein endings of angiosperms is mentioned in 33 publications. Their collective judgment is that sieve tubes may be lacking, may end partway along, or may extend to the vein tip. Most reports conclude that all vein endings are the same within a species, but a few studies mention variation. Only three studies include quantitative data, all based on reconstruction from microtome sections. The present quantitative study, the most extensive one to date, surveyed sieve tubes in vein endings from cleared and stained leaf pieces of Rudbeckia laciniata (Asteraceae, tribe Heliantheae), a North American perennial herb. Of 203 areoles sampled, 32 (15.8%) lacked vein endings whereas 171 (84.2%) had one to six endings per areole. There were 385 individual vein endings in the 171 areoles: 115 lacked sieve tubes, 38 had them to the vein tip, but in most (232) they ended at some intermediate point. Serial cross sections of the latter showed phloem parenchyma continuing beyond the sieve tubes for some distance, beyond which occurred a distal zone consisting solely of tracheary elements. R. laciniata vein endings exhibit the same range of variation reported for dicots in general. More studies combining clearings with selected sections are needed to establish the range of vein ending patterns among angiosperms.     

Quantitative survey of sieve tube distribution in foliar terminal veins of ten dicot species

Quantitative data on sieve tubes in foliar terminal veins (vein endings) were added to the meager published information from only five dicot species. Correlations with other minor vein configurations were also explored. Leaf samples from ten species of dicots (Oxalis nelsonii, O. pes-capri, O. rubra, O. stricta [Oxalidaceae], Caesalpinia pulcherrima. Glycine max, Trifolium repens [Leguminosae], Ampelamus albidus [Asclepidaceae], Eupatorium rugosum [Asteraceae], and Polygonum convolvulus [Polygonaceae]) were selected for two quantitative procedures: 1) a survey of the arrangement of terminal veins and distribution of sieve tubes in terminal veins in 100 areoles per species using stained leaf clearings; and 2) a search for correlations of sieve tube distribution with number and branching patterns of terminal veins, and with sizes of areoles using image analysis. Two Oxalis species (O. pes-capri and O. stricta) had the smallest areoles and virtually no sieve tubes in any terminal vein. Polygonum convolvulus, at the other extreme, had sieve tubes extending to the tips of most terminal veins. The other species had various intermediate sieve tube configurations. The data indicate that species with few or no sieve tubes associated with their terminal veins, regardless of the number of terminal veins per areole, have smaller areoles. These results may have implications regarding the entry of leaf photosynthates into the vascular system.     

贵州小檗属植物(小檗科)叶脉序研究

采用制作叶脉标本和透明叶标本的方法,对贵州产28种2变种小檗属植物叶脉特征进行比较研究。结果表明:贵州小檗属植物的脉序类型有5种:半达缘羽状脉、花环状半达缘羽状脉、简单弓形羽状脉、花环状弓形羽状脉和混合型。叶脉分支一般有五级:1一级脉构架均为羽状脉,粗度有很粗、粗、中等粗细和纤细四种类型,分支方式包括单轴分支和合轴分支;2粗二级脉构架中有分支达缘或分支均不达缘,与中脉夹角变化各异,内二级脉存在或缺失,细二级脉半达缘、真曲行或简单弓形,间二级脉类型复杂多变但频度种间有差异;3三级脉贯串型、结网型或分支型;4四、五级脉网状或自由分支且常混合在一起。脉间区从发育差到良好,小脉从不分支到不均等分支等各种类型均有,叶缘末级脉缺失、不完整、钉状和环状。大部分种类叶缘具齿,每1cm齿数目和齿内腺点的特性等特征在不同种类间有区别,具有鉴定价值,但齿其它特征复杂多变或种间区别较小,同时齿内脉性状也不稳定。此外,齿的有无会对脉序类型产生影响。小檗属植物叶脉类型存在种间差异,具有重要的分类学价值,叶脉类型的变化和复杂程度显示了该属植物的进化特点;叶齿的有无和齿特征具有分类学和系统学意义。基于叶脉特征的研究结果并结合重要的外部形态学特征编制了贵州小檗属植物的分种检索表。研究结果可为小檗属植物分类寻找新的依据并探讨其系统学意义。     

Leaf hydraulic conductance varies with vein anatomy across Arabidopsis thaliana wild‐type and leaf vein mutants

Leaf venation is diverse across plant species and has practical applications from paleobotany to modern agriculture. However, the impact of vein traits on plant performance has not yet been tested in a model system such as Arabidopsis thaliana. Previous studies analysed cotyledons of A. thaliana vein mutants and identified visible differences in their vein systems from the wild type (WT). We measured leaf hydraulic conductance (K_leaf), vein traits, and xylem and mesophyll anatomy for A. thaliana WT (Col‐0) and four vein mutants (dot3‐111 and dot3‐134, and cvp1‐3 and cvp2‐1). Mutant true leaves did not possess the qualitative venation anomalies previously shown in the cotyledons, but varied quantitatively in vein traits and leaf anatomy across genotypes. The WT had significantly higher mean K_leaf. Across all genotypes, there was a strong correlation of K_leaf with traits related to hydraulic conductance across the bundle sheath, as influenced by the number and radial diameter of bundle sheath cells and vein length per area. These findings support the hypothesis that vein traits influence K_leaf, indicating the usefulness of this mutant system for testing theory that was primarily established comparatively across species, and supports a strong role for the bundle sheath in influencing K_leaf.     

Developmental Anatomy of the Leaf of L. Ficus religiosa

The leaves of Ficus religiosa show a more or less sigmoid growthpattern, and under the conditions prevailing in Delhi each leafincreases in size in 9 days from about 425 to 4025 mm²(as judgedby the average mature leaf size) after its emergence from thespathe. The leaf is hypostomatic and has paracytic and anomocyticstomata between polygonal epidermal cells. The frequency ofstomata per mm² increases from 33.3 to 400 and the number oflower epidermal cells decreases from about 11 300 to about 3040per mm² with the growth of the leaves while the number of upperepidermal cells decreases from 5600 to 1110. The vasculaturecomprises a single main vein (the mid rib), secondaries, tertiaries,quaternaries, and intermediaries. The number of areoles permm² decreases from 15.5 to 2.7 while the numbers of vein endingsand vein tips per areole show no correlation either with oneanother or with leaf size.     

HISTOGENESIS OF LEAF VENATION IN TRIFOLIUM WORMSKIOLDII (LEGUMINOSAE)

The histogenesis of the leaf venation in Trifolium wormskioldii was followed, using whole cleared leaflets and transverse and paradermal microtome sections. Mature leaflets are about 8 mm in length. The midvein precambium appears in leaflets 100 μ in length; secondary vein procambium is first seen in 230 μ leaflets; procambium of minor veins first appears in leaflets 400-600 μ in length. Phloem is apparent before xylem, in midvein, secondary veins, and minor veins. Vein endings are initiated and they mature last. No evidence was found to support the theory of “vein breakage.” In the greenhouse 24 days elapsed from leaf initiation to exposure of leaf tips; 6 more days elapsed until full leaf expansion.     

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.     

Quantifying leaf venation patterns: two-dimensional maps

The leaf vasculature plays crucial roles in transport and mechanical support. Understanding how vein patterns develop and what underlies pattern variation between species has many implications from both physiological and evolutionary perspectives. We developed a method for extracting spatial vein pattern data from leaf images, such as vein densities and also the sizes and shapes of the vein reticulations. We used this method to quantify leaf venation patterns of the first rosette leaf of Arabidopsis thaliana throughout a series of developmental stages. In particular, we characterized the size and shape of vein network areoles (loops), which enlarge and are split by new veins as a leaf develops. Pattern parameters varied in time and space. In particular, we observed a distal to proximal gradient in loop shape (length/width ratio) which varied over time, and a margin-to-center gradient in loop sizes. Quantitative analyses of vein patterns at the tissue level provide a two-way link between theoretical models of patterning and molecular experimental work to further explore patterning mechanisms during development. Such analyses could also be used to investigate the effect of environmental factors on vein patterns, or to compare venation patterns from different species for evolutionary studies. The method also provides a framework for gathering and overlaying two-dimensional maps of point, line and surface morphological data.     

中国花楸属单叶类群叶脉序特征研究

中国花楸属单叶类群(24种)包含落萼组与冠萼组,亦有学者将其分为两个属(白花楸属和水榆属)或合并成为一个属(白花楸属).由于其花序和果实的高度相似性,长期以来分类地位具有争议,而叶脉序特征在花楸属单叶类群中的分类作用尚不明确.因此,该研究以国产花楸属落萼组的13个物种和冠萼组的7个物种作为研究对象,以清净法制作标本对叶...     

Sink Effect of Cytokinin in Detached Leaves Is Not Caused by Structural Rearrangements of Phloem

The sink effect of cytokinin is manifested as a decrease in source capacity and the induction of sink activity in the phytohormone-treated region of a mature excised leaf. In order to find out whether this effect was due to the direct action of cytokinin on the phloem structure, two types of phloem terminals were examined. In pumpkin (Cucurbita pepo L.) leaves, the phloem terminals are open; i.e., they are linked to mesophyll by numerous symplastic connections, which are located in narrow areas called plasmodesmal pit fields. In broad bean (Vicia faba L.) leaves, the phloem terminals belong to the closed type and have no symplastic links with mesophyll. The electron microscopic study of terminal phloem did not reveal any structural changes in the companion cells, which could account for the suppression of assimilate export. The treatment of leaves with cytokinin neither disturbed the structure of plasmodesmal pit fields in pumpkin leaves nor eliminated the wall protuberances (the ingrowths promoting phloem loading) in bean leaves. No evidence was obtained that the cytokinin-induced import of assimilates in mature leaves is caused by the recovery of meristematic activity, i.e., by either formation of new phloem terminals having immature sieve elements capable of unloading or by the development of new sieve elements within the existing veins. Cytokinin did not induce de novo formation of phloem elements. Structural characteristics of the leaf phloem, such as the number of branching orders in the venation pattern, the number of vein endings per areole, the number of areoles per leaf, the area of one areole, and the number of sieve elements per bundle remained unaltered. It is concluded that the sink effect of cytokinin in excised leaves cannot be determined by alteration of the phloem structure.     

PARAVEINAL MESOPHYLL IN CALLIANDRA TWEEDII AND C. EMARGINATA (LEGUMINOSAE; MIMOSOIDEAE)

Paraveinal mesophyll (PVM) in Leguminosae, subfamily Mimosoideae, was first reported in 1894 but never described in detail before now. We cleared, and sectioned in resin, leaflets of Calliandra tweedii and C. emarginata (Tribe Ingeae). Lamina anatomy in both species is very similar: one palisade layer, two to three spongy layers, and the horizontal veinal network with its interconnected PVM in between. PVM is a unistratose cellular lacework extending between veins and attached medianly along each flank of all veins. PVM cells have a normal complement of typical chloroplasts similar to other mesophyll cells. Most veins are ensheathed by fibers except for an extended lateral slit along each flank where the PVM is attached; a parenchymatous bundle sheath is therefore lacking. All vein endings lack phloem, although the tracheary elements of some vein endings are flanked by one or two long, slender, seemingly undifferentiated cells. Occasional small gaps occur between the PVM cell wall and adjacent tracheary elements, which expose xylem directly to mesophyll intercellular space. PVM anatomy of Calliandra, including its physical relationship to the various vein orders, differs in some important respects from PVM of the few other leguminous and nonleguminous species studied anatomically in any detail.     

ANASTOMOSES IN THE VENATION OF GINKGO BILOBA

Arnott , Howard J. (Northwestern U., Evanston, Ill.) Anastomoses in the venation of Ginkgo biloba. Amer. Jour. Bot. 46(6): 405–411. Illus. 1959.—Although the vasculature of the lamina of Ginkgo biloba has been described as open dichotomous and devoid of anastomoses, vein unions were found in a survey of 1065 leaves collected from both long and short shoots of 11 trees. When studied by directing a strong light through the lamina and by clearings, 9.9% of the leaves possessed 1 or more anastomoses. Long shoot-leaves showed 13.4% anastomoses while short shoot-leaves showed 8.2% anastomoses. Multiple anastomoses were found in almost half of the leaves bearing anastomoses. In the 105 leaves having vein unions, 163 anastomoses were counted. These anastomoses could be grouped into 4 types which are easily distinguished by the number of dichotomies involved and by the presence or absence of dichotomies above the point of vein union. Other deviations were found from the “normal” venation pattern; these consisted of unconnected veins, veins anastomosed marginally but unconnected basally, and veins ending a considerable distance from the margin. It was speculated that the anastomoses found in Ginkgo biloba are of a simple, archaic type and are apparently analogous to the anastomoses in the leaves of certain ferns and in the leaflets of various cycads. The evolutionary significance of these anastomoses must be assessed by a broad study of venation patterns in the seed-plants.     

中国连香树科的系统研究——Ⅰ.叶的宏观结构及叶柄维管束变化

本文研究了连香树科(Cercidiphyllaceae)叶的宏观结构,首次报道连香树齿腺体显微结构及晶体类型,并对叶柄维管束的变化作了进一步研究。通过与近缘科的比较,我们认为连香树科的系统演化处于孤立地位,和金缕梅科有较近的亲缘关系,与木兰科较为疏远。     

Tracheid analysis and modeling of the minor veins of the coleus and smilax leaves

Tracheid analysis was carried out on the veinlets and minor veins of the coleus (Solenostemon scutellarioides [L.] Codd) leaf. Third- to fifth-order, or minor, veins average 3.4 tracheids in tandem and they bipartition islets when these enclosed islets reach a critical size; both these features of vein length and islet size contribute to a self-similar process of vein pattern generation. An areole was calculated to be initially comprised of about ten cells making the patterning event for vein formation requiring only a few cells. An algorithmic model developed here for minor vein formation includes five production rules, and this computer model explains the 3–4 tracheids per minor vein, presence of isolated tracheids, the structure of veinlets, and the elaborate branching patterns of veinlets in coleus and other plants.