Studies on the leaf anatomy of Euphorbia: II. Venation patterns*

Leaf venation patterns of 150 species of Euphorbia are presented and their value as a diagnostic feature in the genus assessed. Based on the gross venation patterns, the species have been grouped into uni-, bi-, tri-veined and special categories. The majority of the species studied belonged to the tri-veined category, in which ornamentation of the veins and the course of traces in the lamina proved useful additional characters. Features such as the size of the areoles, the number of vein endings, and their further ramifications and composition in each areole varied in the same leaf or in different leaves of a given species. Furthermore, no direct correlation could be established between the areole size and the numbers of vein endings and tips per areole. Forty species possess a parenchymatous vein sheath. Globular chloroplasts showed up conspicuously in the sheath cells of a few species. Dilated tracheidal elements, localized on the venules or at the tips of vein endings, are characteristic of the xerophytic species. In some instances, correlation was apparent between plants grouped according to their venation patterns and their habit.     

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.     

滇中半湿润常绿阔叶林主要优势及常见种叶形态结构特征

通过对滇中亚热带半湿润常绿阔叶林８２个主要优势种和常见种进行叶结构分析研究,结果表明这些主要组成树种虽亲缘关系相互较远,但在相同的生态环境条件下叶结构却表现出了相似和相同的特点。主要为：群落中以单叶为主,边缘具齿较全缘为多,缺长渐尖叶,有毛叶和无毛叶所占的比例近于相等;叶脉主要为环结曲行羽状脉,大部分二级脉结环,被三、四级脉环所包围,夹角为锐角且变异多近一致;三级脉多为结网型;网眼发育完善较不完善为多,盲脉多分枝;大部分无包藏脉。并且讨论了其中一些特征如毛被,网眼,包藏脉和边缘脉等可能的生态学意义。该类森林群落主要树种的叶形态结构特征除了是物种自身系统进化的原因外,还是植物长期适应滇中高原偏干旱的季风气候的表现,是生态适应在叶形态结构上的反映     

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.     

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.     

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.     

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.     

中国木犀属植物叶脉形态及其分类学意义

观察了中国木犀属植物4组19种叶脉形态。主要分析木犀属植物叶片脉序走向,脉序为环结曲行或半直行羽状脉。二级脉急转曲行或半直行,叶脉分支一般为4级,少数5级。盲脉1～2次分支,少数3次或不分支。仅柊树叶缘末级脉汇合成边脉.部分叶缘具齿,叶缘齿性状不稳定,因其内主脉不同而在本属种间表现出一个连续的变异过程。圆锥花序组与李榄属和木犀榄属从叶片脉序特征方面表现出较近的亲缘关系。四个组的叶脉形态在演化上关系上与花粉形态表现相一致。编写了叶片脉序特征分种检索表。     

Leaf architecture and anatomy of eleven species of Mortoniodendron (Malvaceae s.l.)

The genus Mortoniodendron is distributed from southeastern Mexico to Colombia and contains about 18 species of trees and shrubs. The main goal of this study is to describe and compare the leaf architecture and the anatomy of 11 species of Mortoniodendron using cleared leaves and paradermal and transverse leaf sections. All of the species of Mortoniodendron share a simple lamina with an entire margin. The differences among the species include the venation pattern, the relative thickness and course of the secondary veins, the presence or absence of the intersecondary veins, the tertiary vein pattern, the number of branches in veinlets, and the marginal ultimate venation. The leaf anatomy is similar to other species of the Malvaceae s.l.: for example, mucilage cells are common in Mortoniodendron. There are unique attributes of the leaf architecture and anatomy, such as the eucamptodromous venation pattern of M. guatemalense, which facilitates the identification of the species. The species M. palaciosii, M. pentagonum, M. ruizii, M. uxpanapense, and M. sp. nov. share features that suggest that they are closely related.     

Foliar architecture and anatomy of <Emphasis Type="Italic">Bernardia</Emphasis> and other genera of Acalyphoideae (Euphorbiaceae)

A comparative study of leaf architecture and anatomy of 42 species of Bernardia and other genera related of Acalyphoideae was undertaken to identify characters that support infrageneric and specific delimitation. Thirty variable foliar architectural and anatomical characters were studied. Some characters are consistent (e.g., venation pattern, secondary and tertiary vein arrangement, presence or absence of agrophic veins, type of areoles, bundle sheath cell type, presence or absence of fibers in the mesophyll, trichome type, stomata location, and type of crystals), and characterize genera. In addition, foliar character distribution within Bernardia supports the most recent infrageneric classification.     

Leaf Venation and Its Systematic Significance in Sapindaceae of China

Leaf venation of 27 species representing 25 genera of Sapindaceae (sstr.) of China was investigated for the first time. The pinnate venation pattern in most species is either camptodromous, or craspedodromous. Three types of leaf blade margin were observed, ie., entire, toothed and partite. The secondary veins are branched or unbranched. Most species have intersecondary veins. The tertiary veins of most species are reticulate and percurrent. The areoles are regular or irregular. Veinlets are simple, branched or absent. The delimitations of Xanthoceroideae and Lepisanthes sensu lato are supported by leaf venation characters. The close relationships among Dimocarpus, Litchi and Nephelium are supported by the evidence from leaf venation. A key to the species of Sapinaceae based on leaf venation characters is presented.     

中国无患子科植物的叶脉形态及其系统学意义

对国产狭义无患子科25属27种植物的叶脉形态特征进行了研究报道。结果表明：叶脉均属于羽状脉类型,其中多数为曲行羽状脉,部分为直行羽状脉;叶缘有全缘、具齿和深裂3种类型;二级叶脉具有分支和不分支两种类型;大部分种类具二级间脉,少数不具间脉或间脉不明显;多数种类的三级脉为结网型和贯串型并存;网眼的发育有完善和不完善2种类型;盲脉有简单、具分支和无盲脉3种类型。叶脉形态研究结果支持文冠果亚科以及广义鳞花木属概念,观察发现龙眼属、荔枝属与韶子属从脉序特征方面表现出较近的亲缘关系。编写了国产无患子科叶片脉序特征检索表。     

国产柳属繁柳组植物叶脉序研究及其分类学意义

采取脉络法,对柳属（Salix）繁柳组（Sect.Denticulatae C.K.Schneid.）17种及1变种进行叶片脉序观察。结果表明：繁柳组植物叶脉序类型均为直行羽状脉,表现出较高的一致性。该组植物均有二级间脉,但类型各有不同;三级脉及较高级脉序大体有三种类型;网眼发育完善或不完善;盲脉偶有不分枝;不同种之间网眼的发育以及盲脉分枝具有显著差异,是有价值的分类性状。根据聚类分析,繁柳组植物叶脉序分为两种类型：草地柳（S.praticola Hand.-Mazz.ex Enander）类型及腹毛柳（S.delavayana Hand.-Mazz.）类型,研究结果与形态学结果基本一致,但叶脉序提供了新的分类学性状,可为柳属植物疑难种的分类寻找新的依据。     

Fossil Nelumbonaceae from the La Colonia Formation (Campanian–Maastrichtian,Upper Cretaceous), Chubut,Patagonia, Argentina

We describe here fossil leaves and fruits assignable to the family Nelumbonaceae Dumortier 1828. Fossils were collected at the Cañadón del Irupé locality, La Colonia Formation (Campanian–Maastrichtian, Upper Cretaceous), Chubut Province, Patagonia, Argentina. The fossil leaves are simple, with symmetrical and peltate lamina, orbicular in shape, and an entire margin. The venation is truly actinodromous with at least 15 primary veins, the secondary veins are poorly developed and intercalated with the primary veins, the third category vein is opposite, and the areoles are well-developed and 4- to 5-sided. These features are characteristic of the extant genus Nelumbo Adamson 1763, and the presence of these characters in the fossils allows their placement within this genus. Reproductive structures, similar to the fruit-receptacle of Nelumbo with fruits in situ, were found associated but not in organic connection with the leaves. These fossils constitute the only and oldest record for the family in the Southern Hemisphere. This record indicates that Nelumbo was more widespread in the past than today.     

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。     

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.     

CLASSIFICATION OF THE ARCHITECTURE OF DICOTYLEDONOUS LEAVES

A classification of the architectural features of dicot leaves—i.e., the placement and form of those elements constituting the outward expression of leaf structure, including shape, marginal configuration, venation, and gland position—has been developed as the result of an extensive survey of both living and fossil leaves. This system partially incorporates modifications of two earlier classifications: that of Turrill for leaf shape and that of Von Ettingshausen for venation pattern. After categorization of such features as shape of the whole leaf and of the apex and base, leaves are separated into a number of classes depending on the course of their principal venation. Identification of order of venation, which is fundamental to the application of the classification, is determined by size of a vein at its point of origin and to a lesser extent by its behavior in relation to that of other orders. The classification concludes by describing features of the areoles, i.e., the smallest areas of leaf tissue surrounded by veins which form a contiguous field over most of the leaf. Because most taxa of dicots possess consistent patterns of leaf architecture, this rigorous method of describing the features of leaves is of immediate usefulness in both modern and fossil taxonomic studies. In addition, as a result of this method, it is anticipated that leaves will play an increasingly important part in phylogenetic and ecological studies.     

Linking leaf veins to growth and mortality rates: an example from a subtropical tree community

A fundamental goal in ecology is to link variation in species function to performance, but functional trait–performance investigations have had mixed success. This indicates that less commonly measured functional traits may more clearly elucidate trait–performance relationships. Despite the potential importance of leaf vein traits, which are expected to be related to resource delivery rates and photosynthetic capacity, there are few studies, which examine associations between these traits and demographic performance in communities. Here, we examined the associations between species traits including leaf venation traits and demographic rates (Relative Growth Rate, RGR and mortality) as well as the spatial distributions of traits along soil environment for 54 co‐occurring species in a subtropical forest. Size‐related changes in demographic rates were estimated using a hierarchical Bayesian approach. Next, Kendall's rank correlations were quantified between traits and estimated demographic rates at a given size and between traits and species‐average soil environment. Species with denser venation, smaller areoles, less succulent, or thinner leaves showed higher RGR for a wide range of size classes. Species with leaves of denser veins, larger area, cheaper construction costs or thinner, or low‐density wood were associated with high mortality rates only in small size classes. Lastly, contrary to our expectations, acquisitive traits were not related to resource‐rich edaphic conditions. This study shows that leaf vein traits are weakly, but significantly related to tree demographic performance together with other species traits. Because leaf traits associated with an acquisitive strategy such as denser venation, less succulence, and thinner leaves showed higher growth rate, but similar leaf traits were not associated with mortality, different pathways may shape species growth and survival. This study suggests that we are still not measuring some of key traits related to resource‐use strategies, which dictate the demography and distributions of species.     

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.