基于核糖体DNA内转录间隔区ITS序列的广义蓼属及近缘属分子系统学研究(英文)

通过对广义蓼属及近缘属共32个代表种内转录间隔区ITS序列的分子系统学分析,尝试研究备受争议的广义蓼属及近缘属的物种族、属、组级的划分问题,结果显示,广义蓼属在系统发育树上并不能形成一个单系类群,这些物种共聚为3大支,分别对应春蓼族、蓼族及荞麦族,其中荞麦属与翅果蓼属形成了一支独立于春蓼族及蓼族之外的类群。在春蓼族中,冰岛蓼属与分叉蓼组形成一个单系类群。     

基于核糖体DNA内转录间隔区ITS序列的广义蓼属及近缘属分子系统学研究

通过对广义蓼属及近缘属共32个代表种内转录间隔区ITS序列的分子系统学分析,尝试研究备受争议的广义蓼属及近缘属的物种族、属、组级的划分问题,结果显示,广义蓼属在系统发育树上并不能形成一个单系类群,这些物种共聚为3大支,分别对应春蓼族、蓼族及荞麦族,其中荞麦属与翅果蓼属形成了一支独立于春蓼族及蓼族之外的类群。在春蓼族中,冰岛蓼属与分叉蓼组形成一个单系类群。     

Generic limits in Polygonurn and related genera (Polygonaceae) on the basis of floral characters

In order to investigate the taxonomic value of a number of characters in the elucidation of relationships within the collective genus Polygonum L., and segregate genera Atraphaxis L., Calligonum L., Pteropyrum Jaub. & Spach., Oxygonum Burch., Fagopyrum Mill., Harpagocarpus Hutch. & Dandy and Polygonella Michx., 83 species of the tribe Polygoneae emend. Jaretzky were studied. The following floral characters were examined: stamen number, insertion and external morphology; structure and morphology of the floral nectaries; vasculature and surface morphology of the tepals.
Problems in floral morphology in Polygonaceae are discussed, together with relationships within the Family. The tribal division proposed by Haraldson (1978) is supported, but a new and different delimitation of genera is proposed. The present data support a division of Polygonum L. sensu lato and its related taxa into two tribes: Polygoneae (containing Polygonum L. sensu stricto, Fallopia Adans., Oxygonum Burch., Pteropyrum Jaub. & Sparh., Atraphaxis L. and Calligonum L.), and Persicarieae (containing Fagopyrum Mill., Harpagocarpus Hutch. & Dandy, Persicaria Mill. and Koenigia L.). The genus Reynoutria Houtt. is included as a section of Fallopia Adans.; Bislorta Mill., Aconogonon Reichenb. and Tovara Adans. are included within Persicaria Mill.
A number of new combinations are proposed in Persicaria and Fallopia .     

29种中国野豌豆属植物叶表皮微形态特征及其系统学意义

应用光学显微镜和扫描电子显微镜对29种野豌豆属植物叶表皮微形态特征进行了观察。结果显示:叶表皮细胞形状有无规则形和多边形2种,垂周壁式样有深波状、浅波状和平直-弓形;表皮角质层纹饰微形态多样,大多数植物叶片表面不具有腺毛或仅中脉有,少数植物叶片表面具腺毛;部分叶表皮上有柔毛,少数植物无毛。气孔器存在于上表皮、下表皮、或上下表皮均有,形状为椭圆形、卵圆形,均为无规则型。野豌豆属植物叶表皮的这些微形态特征,在属内组间没有明确的规律性,但可为探讨该属种间的分类学及亲缘关系提供依据。     

Reappraisal of the generic status of Pteroxygonum (Polygonaceae) on the basis of morphology,anatomy and nrDNA ITS sequence analysi

Gross morphology, fruit anatomy, tepal venation, pollen morphology, chromosome number and ITS sequence of Pteroxygonum Damm. & Diels as well as other related genera (Polygonum, Fallopia, Reynoutria, Fagopyrum, and Antenoron) have been investigated to evaluate the generic status of Pteroxygonum. Pt. giraldii Damm. & Diels has three sharp horns at the base of fruit, which is distinctive among all the genera investigated. Upon observation of fruits under a light microscope (LM), the exocarp of Pt. giraldii is usually thickened and delimited by the rectangular cells with some sporadic undulating lumen, while that of Fagopyrum is thin-walled and isodiametric to rectangular in the cell shape. Analysis of tepal venation was performed under a stereomicroscope, and two types of tepal venation were found in Fagopyrum and Pteroxygonum. The type I is trifid, observed in Pt. giraldii, F. esculentum Moench, F. dibotrys (D. Don) Hara and F. tataricum (L.) Gaertn. The type II, found in F. caudatum (Sam.) A. J. Li, F. urophyllum (Bur. & Franch.) H. Gross and F. gracilipes (Hemsl.) Damm. ex Diels, has the main vein extending from tepal base with some secondary veins. Evidence from tepal venation supports the previous classification in which Fagopyrum can be divided into a large-achene group and a small-achene group. Pollen morphology was investigated under a scanning electron microscope (SEM). The exine ornamentation of Pt. giraldii was finely reticulate with lumina diameter wider than muri width. The exine ornamentation in all the examined Fagopyrum species is, however, prominently sunken punctuate. The phylogenetic analysis of nuclear ribosomal DNA (nrDNA) ITS sequences in Pteroxygonum and related genera indicated that all the species form a well-supported monophyletic group with two clades. One includes Polygonum sect. Avicularia Meisn., genus Fallopia and genus Reynoutria, and the other consists of other sections of Polygonum, genus Fagopyrum and Pteroxygonum. The latter clade can be divided into two subclades. Fagopyrum species compose the first one, while Pteroxygonum giraldii, species of Polygonum (except sect. Avicularia) and Antenoron form the second one. In consideration of the above evidence, we conclude that Pteroxygonum is an independent genus in tribe Persicarieae, and should not be merged into the genus Fagopyrum.     

对翼蓼(蓼科)属级位置的重新探讨

通过对翼蓼Pteroxygonum giraldii Damm. &; Diels及相关属(蓼属Polygonum、何首乌属Fallopia、虎杖属Reynoutria、荞麦属Fagopyrum和金线草属Antenoron)的形态观察、果实解剖学观察、花被片脉序观察、花粉形态、核型分析, 以及ITS序列的分析确定了翼蓼和荞麦F. esculentum Moench较远的亲缘关系。其中我们发现翼蓼果实基部有三个角状物明显不同于其他属果实的形态特征。翼蓼外果皮明显加厚, 并有零星散布的波状内腔, 而荞麦的外果皮很薄, 细胞不等径, 中果皮极厚。以上证据证明了翼蓼与荞麦属亲缘关系较远。在观察荞麦属和翼蓼的花被片脉络时发现了两种不同的脉序类型, 符合将荞麦属分为两个组的划分。翼蓼花被片脉序为三出状, 支持将翼蓼归为Persicarieae族。对翼蓼及荞麦属植物的花粉进行比较后, 发现荞麦属植物的花粉网孔有明显的内凹穿孔而翼蓼却没有, 结果表明二者亲缘关系较远。通过对nrDNA ITS区域序列分析得出翼蓼及相关属为一个单系类群, 含有两个稳定的分支: 第一个分支由蓼属(萹蓄组sect. Avicularia)、何首乌属、虎杖属的植物组成, 第二个分支由蓼属(刺蓼组sect. Echinocaulon、蓼组sect. Polygonum、分叉蓼组sect. Aconogonon、拳参组sect. Bistorta、翼蓼和荞麦属植物组成。同时第二个分支又分成了两个亚分支, 蓼属(刺蓼组、蓼组、分叉蓼组、拳参组)和翼蓼属Pteroxygonum植物属于第一个亚支而荞麦属植物属于第二个亚支。结果支持翼蓼不属于荞麦属的范畴。实验结果显示翼蓼是个单型属, 属于Persicarieae族。     

十字花科四属植物叶片的表皮特征

对十字花科岩荠属、阴山荠属、泡果荠属、棒毛荠 属植物的叶表皮特征进行了扫描电镜观察,结果表明：表皮细胞多为不规则形,气孔主要分布于表皮,垂周壁上有明显的波状嵴或沟槽状下陷,平周壁上有各咱条纹状角质层纹饰。这些微形态特征在属间有明显的类群特异性,又有一定的过渡类型,为岩荠属及近缘属的合理划分提供一定的证据。     

Characters of Leaf Epidermis and Their Systematic Significance in Menispermaceae

The epidermal characters of mature leaves of 29 genera, 50 species and 3 varieties (totally 56 samples) representing all the 5 tribes in the family Menispermaceae were examined under the light microscope. The main conclusions are as follows: (1) The shape of the epidermal cells is polygonal or irregular, and the anticlinal walls are straight or waved in the family. In some genera a special arrangement of epidermal cells is named as “rosette-cell arrangement" for the first time. The lower epidermal cells were found to have a papilla in Stephania, Diploclisia and Legnephora . (2) In some genera, the anticlinal walls are oblique, rather than perpendicular, to the surface. (3) The stomatal apparatuses, generally restricted to the lower surface of the leaves, were assigned to anomocytic, staurocytic, cyclocytic, anisocytic and actinocytic types, and their distribution on the epidermis may be of diffuse pattern or island congregating pattern. (4) The cells of both upper and lower epidermis in the tribe Pachygoneae and Fibraureae are generally polygonal with straight or arched anticlinal walls, and the stomatal apparatuses are usually staurocytic and actinocytic. The cyclocytic stomatal apparatus was found only in two genera of the tribe Pachygoneae. By contrast, the epidermal cells of the tribes Anomospermeae, Tinosporeae and Menispermeae are generally irregular with waved anticlinal walls, and the stomatal apparatuses are predominantly anomocytic. These correlated characters are of much significance in delimiting tribes within the Menispermaceae, and also provide evidence for studies on systematic relationships of several genera.     

Fine structural observations on the epidermis

Summary The organization of the wall of epidermal cells in the petiole of species of Apium, Eryngium, Rumex, and Abutilon as well as that of the epidermis of Avena coleoptile has been investigated. The outer and inner tangential walls consist of layers in which the cellulose microfibrils are oriented alternately parallel or transverse to the longitudinal cell axis. This organization resembles that previously described for collenchyma cell walls (Wardrop, 1969; Chafe, 1970). On the radial (anticlinal) walls the orientation of the microfibrils is transverse and these appear continuous with the layers of transverse orientation of the outer and inner tangential walls. Variation in thickness of the outer tangential, and radial, and inner tangential walls appears to result from the variation in thickness of those layers in which the microfibrils have a longitudinal orientation. The extent to which these observations can interpreted in terms of some type of modified multi-net growth is discussed.     

Seed testa micromorphology of thirty‐eight species of Allium (Amaryllidaceae) from central Asia,and its taxonomic implications

Micromorphological characteristics of the seed testa of 38 species belonging to 19 sections and seven subgenera of Allium from central Asia have been evaluated with a scanning electron microscope (SEM). Several taxonomically significant characters were found. The results showed that the epidermis consists of polygonal cells with different dimensions, that the cellular arrangement is tight or loose with or without reticulate tissue, and that the arrangement of cells is regular or irregular. The main variation was found in the anticlinal walls, of which the outline could be straight, straight to arched, arched or undulating like an S or U. Convex and (or) concave verrucate periclinal walls occurred in all species investigated. Based on the characteristics of the anticlinal and periclinal walls of the 38 species, six groups of Allium were distinguished, and a dichotomous key for them is presented. The cellular arrangement and periclinal and anticlinal wall traits of the 38 species of Allium were found stable and distinct within a species, but showed great differences among species. We conclude that seed epidermal characteristics provide useful and important information for distinguishing species of Allium, and thus they have taxonomic significance.     

Biomacromolecules of Algae and Plants and their Fossil Analogues

A review of our current understanding of resistant biomacromolecules derived from present and past algae and higher plants is presented. Insight in the nature of recent and fossil macromolecules is strongly hampered by the difficulties in obtaining the material in pure and unaltered form. For the extant material, avoiding artificial condensation and structural alteration as a result of chemical isolation and purification of biomacromolecules requires constant attention. To date, several types of sporopollenin seem to occur. One type is characterised by oxygenated aromatic building blocks, in particular p-coumaric acid and ferrulic acid. The other type is thought to consist predominantly of an aliphatic biopolymer. In this review it is concluded that extant sporopollenin consists of the aromatic type, whereas the aliphatic component of fossil sporopollenin is due to early-diagenetic oxidative polymerization of unsaturated lipids. The cuticles of most higher plants contain the aliphatic biopolyester cutin. Additionally, cuticles of drought-adapted, mostly CAM plants, seem to contain the non-hydrolysable aliphatic biopolymer cutan. Only a very few algae are able to biosynthesize resistant, (fossilisable) cell walls: some Chlorophyta, Eustigmatophyta and Prasinophyta produce the aliphatic biopolymer algaenan. Some Dinophyta are also capable of producing algaenan cell walls. Additionally, some taxa produce highly resistant cyst-walls with a high proportion of aromatic moieties. For the morphologically well-preserved fossil material, contamination by organic particles other than the target taxon is hard to eliminate and can contribute to either the aliphatic or aromatic signal. Furthermore, post-mortem migration of aliphatic moieties into, and their condensation onto the macromolecule might occur, e.g. by oxidative polymerization. These phenomena hamper the evaluation of the aliphatic signature of fossil plant material and may for example explain the preservation of initially cutin-based cuticles or non-algaenan containing algae. The extent to which migration and in situ formation of aromatic moieties plays a role in modifying resistant algal macromolecules, notably under elevated temperature and/or pressure conditions, still remains an open question.     

Pollen morphology of the genera Polygonum s. str. and Polygonella (Polygoneae: Polygonaceae)

The pollen of 30 taxa (27 species, one subspecies and two varieties) in two genera, viz Polygonum s. str. and Polygonella was investigated with LM and SEM, and some selected taxa with TEM. In all genera investigated the pollen is prolate to spheroidal, and the aperture is mostly tricolporate, rarely panto-hexacolporate (especially Polygonum section Polygonum). The exine sculpturing pattern is the most variable feature. Three types of exine can be recognized. Type 1 (Avicularia-Type, sensu Hedberg) - All species of section Polygonum and section Tephis share the smooth tectate exine with spinules, sometimes the surface is more or less rough (Polygonum afromontanum in section Tephis). Type 2 (Pseudomollia-Type, sensu Hong) - Pollen of Polygonum molliaeforme (section Pseudomollia) has the exine, which is verrucose on both poles and nearby the mesocolpium, and mostly psilate around the ectoaperture. Type 3 (Duravia-Type, sensu Hedberg) - Pollen grains of Polygonum section Duravia and Polygonella have the exine which is semitectate-reticulate at the mesocolpium and the poles, and rugulate/reticulate or sometimes foveolate with microspinules around the ectoapertures. The pollen grains in four taxa (viz Polygonum section Pseudomollia, P. section Duravia and genus Polygonella) have a well-marked dimorphism of the ektexine, which is considered to be a synapomorphic condition. The differences of pollen grain between the genus Polygonella and Polygonum section Duravia are almost non existent and clearly interrelated. It is therefore postulated that the similarity in pollen of both taxa is not the result of convergency, but is interpreted as a homology. It is noteworthy that the pollen of Polygonum molliaeforme (section Pseudomollia) appears as intermediate between the Avicularia-type and the Duravia-type, and is well supported the value of separated section for its own. Additionally, in TEM, some exine ultrastructures (e.g. columellae, foot layer, endexine) appear to be valuable characters for comparison between/among taxa. The systematic potentialities of the pollen data of the studied taxa at various systematic levels are also discussed.     

中国蓼属头状蓼组植物叶表皮微形态及其分类学意义

采用光学显微镜对中国蓼属头状蓼组17种7变种植物的叶片下表皮微形态进行了观察研究,结果表明,其叶片下表皮微形态特征分为4种类型:(1)气孔器类型为无规则型,表皮细胞不规则形,垂周壁波状或深波状;(2)非典型不等型,偶有无规则型,表皮细胞多边形或不规则形,垂周壁弓形、波状或深波状;(3)平列型,表皮细胞不规则形,垂周壁深波状;(4)平列型兼有非典型不等型,表皮细胞不规则形,垂周壁波状。根据其叶片下表皮气孔器类型,结合该组植物形态、习性等特征,将中国蓼属头状蓼组植物划分为4个系,即掌裂叶系、多年生系、蓼子草系以及一年生直立系。     

Seed micro‐morphology and its systematic significance in Gypsophila (Caryophyllaceae) and allied genera

In order to examine the systematic application of seed‐coat micro‐morphology in Gypsophila and allied genera, seed surfaces of 30 species and two varieties representing four genera of tribe Caryophylleae were examined with scanning electron microscope. The seeds of examined species range between 0.5–2.1 mm in length and 0.3–2.4 mm in diameter. The exomorphology of the seed coat shows two distinctive cell patterns. The epidermis is constructed either of elongated polygonal or of broad polygonal cells. The elongated type is the most common among the studied species, but the variation in alignment of testa cells, their size and shape as well as the density of protuberances may provide further information and useful diagnostic characters at generic and specific rank. The testa cells in Gypsophila and Saponaria are shallowly undulate, deeply undulate, lobed and armed at anticlinal walls. Deeply undulate anticlinal walls were observed in both Gypsophila and Ankyropetalum and a few species of Saponaria. Non‐ or indistinctly grooved anticlinal walls is the more common type in Allochrusa. Seed‐coat characters support the separation of Gypsophila and Saponaria to some extent but disagree with recognition of Ankyropetalum as a genus separate from Gypsophila.     

Rearrangements of F-actin during Stomatogenesis Visualised by Confocal Microscopy in Fixed and Permeabilised Tradescantia Leaf Epidermis

Abstract: New details of F-actin organisation in leaf epidermal and stomatal cells were revealed by rhodamine — and fluorescein — phalloidin staining of fixed epidermal peels of Tradescantia virginiana and visualisation by confocal microscopy. Non-specialised epidermal cells contain highly organised arrays of fine cortical actin filaments aligned in transverse or oblique orientations. In interphase guard mother cells (GMCs), the arrangement of cortical F-actin changes on the periclinal and anticlinal cell walls at different times during differentiation. Initially, cortical F-actin on the periclinal surfaces is oriented transversely and F-actin is evenly distributed around the anticlinal walls. Following polarisation of the adjacent subsidiary mother cells (SMCs), actin in GMCs concentrates on the lateral anticlinal walls, but not on the transverse walls. Subsequently, F-actin on the periclinal walls reorients to radial and then longitudinal. Organisation of F-actin in SMCs appears to be influenced by the adjacent GMCs and co-ordination in F-actin arrangements in cells of the stomatal complex continues through to the formation of the guard cell pair. Our studies indicate that actin bands marking the division site in prophase cells, and detected in microinjected living material, are a particularly labile subset of F-actin. Actin bands were difficult to preserve, even when aldehyde fixation was avoided, in contrast to all interphase and mitotic F-actin.     

国产野豌豆属植物叶表皮微形态特征及其系统学意义

应用光学显微镜和扫描电子显微镜对29种野豌豆属植物叶表皮微形态特征进行了观察。结果显示：叶表皮细胞形状有无规则形和多边形2种,垂周壁式样有深波状、浅波状和平直-弓形;表皮角质层纹饰微形态多样,大多数植物叶片表面不具有腺毛或仅中脉有,少数植物叶片表面具腺毛;部分叶表皮上有柔毛,少数植物无毛。气孔器存在于上表皮、下表皮、或上下表皮均有,形状为椭圆形、卵圆形,均为无规则型。野豌豆属植物叶表皮的这些微形态特征,在属内组间没有明确的规律性,但可为探讨本属种间的分类学及亲缘关系提供依据。     

Cell wall polysaccharides from fern leaves: evidence for a mannan-rich Type III cell wall in Adiantum raddianum

Primary cell walls from plants are composites of cellulose tethered by cross-linking glycans and embedded in a matrix of pectins. Cell wall composition varies between plant species, reflecting in some instances the evolutionary distance between them. In this work the monosaccharide compositions of isolated primary cell walls of nine fern species and one lycophyte were characterized and compared with those from Equisetum and an angiosperm dicot. The relatively high abundance of mannose in these plants suggests that mannans may constitute the major cross-linking glycan in the primary walls of pteridophytes and lycophytes. Pectin-related polysaccharides contained mostly rhamnose and uronic acids, indicating the presence of rhamnogalacturonan I highly substituted with galactose and arabinose. Structural and fine-structural analyses of the hemicellulose fraction of leaves of Adiantum raddianum confirmed this hypothesis. Linkage analysis showed that the mannan contains mostly 4-Man with very little 4,6-Man, indicating a low percentage of branching with galactose. Treatment of the mannan-rich fractions with endo-β-mannanase produced characteristic mannan oligosaccharides. Minor amounts of xyloglucan and xylans were also detected. These data and those of others suggest that all vascular plants contain xyloglucans, arabinoxylans, and (gluco)mannans, but in different proportions that define cell wall types. Whereas xyloglucan and pectin-rich walls define Type I walls of dicots and many monocots, arabinoxylans and lower proportion of pectin define the Type II walls of commelinoid monocots. The mannan-rich primary walls with low pectins of many ferns and a lycopod indicate a fundamentally different wall type among land plants, the Type III wall.     

Epidermal anatomy of Barthelopteris germarii from the Upper Carboniferous and Lower Permian of France and Germany

The epidermal anatomy of Barthelopteris germarii, a late Paleozoic seed fern, is described on the basis of material from the Upper Stephanian of Central France. A number of features are described for the first time for this species. Some have never been reported for late Paleozoic pteridosperms. Although our material is in most respects very similar to previously described material of B. germarii from Germany, there are also some striking differences, especially the peltate glandular trichomes that are very common in the French but completely absent in the German material. Their presence is most probably an ecological adaptation as in many modern plants. This and other epidermal and gross-morphological features (e.g., the presence of papillae on subsidiary cells of the stomata, anastomosing venation) indicate that B. germarii was well adapted to stressed conditions. Of more general interest is the preservation of the material. There appears to exist a clear relationship between the preservation of anticlinal walls and the thickness of the parenchymatic mesophyll; anticlinal walls are very well preserved where parenchyma was thin. Therefore, the absence of anticlinal walls in fossil cuticles, which has often been used as a taxonomic character, is not necessarily a primary feature.     

In situ analysis by microspectroscopy reveals triterpenoid compositional patterns within leaf cuticles of Prunus laurocerasus

The cuticular waxes on the leaves of Prunus laurocerasus are arranged in distinct layers differing in triterpenoid concentrations (Jetter et al., Plant Cell Environ 23:619–628, 2000). In addition to this transversal gradient, the lateral distribution of cuticular triterpenoids must be investigated to fully describe the spatial distribution of wax components on the leaf surfaces. In the present investigation, near infrared (NIR) Raman microspectroscopy, coherent anti-Stokes Raman scattering (CARS) microscopy, and third harmonic generation (THG) spectroscopy were employed to map the triterpenoid distribution in isolated cuticles from adaxial and abaxial sides of P. laurocerasus leaves. The relative concentrations of ursolic acid and oleanolic acid were calculated by treating the cuticle spectra as linear combinations of reference spectra from the major compounds found in the wax. Raman maps of the adaxial cuticle showed that the triterpenoids accumulate to relatively high concentrations over the periclinal regions of the pavement cells, while the very long chain aliphatic wax constituents are distributed fairly evenly across the entire adaxial cuticle. In the analysis of the abaxial cuticles, the triterpenoids were found to accumulate in greater amounts over the guard cells relative to the pavement cells. The very long chain aliphatic compounds accumulated in the cuticle above the anticlinal cell walls of the pavement cells, and were found at low concentrations above the periclinals and the guard cells.