夹竹桃科植物叶表皮特征及其系统学研究

以萝藦科植物匙羹藤为对照,在扫描电镜下观察了传统上分属于夹竹桃科3亚科植物的叶下表皮特征。结果表明:(1)所观察的夹竹桃科25种植物叶下表皮特征具有一定程度的多样性,如少数物种具有表皮毛;部分角质层表面有丝状蜡质;叶下表皮纹饰有条纹状、脊状皱褶增厚、块状突起、花蕊形和较平滑等类型;气孔大小差异较大,气孔外拱盖和内缘形状分浅波状和较平滑2种类型,气孔保卫细胞极区有T型加厚和无T型加厚两种类型,气孔外拱盖与表皮细胞间分凹陷和连续两种类型等,这些叶表皮特征对夹竹桃科植物属内和属间类型的分类以及对疑难类群的划分具有重要的参考价值。(2)夹竹桃科25种植物的绝大部分种的叶表皮气孔类型、气孔外拱盖及其内缘形态基本一致,但叶表皮纹饰、气孔外缘角质层特征呈现出较高的多样性,在属内各种和属间有较大的差异。(3)萝藦科植物匙羹藤与夹竹桃科植物盆架树在气孔形状,叶片气孔外缘角质层特征、气孔外拱盖等叶表皮特征具有高度的相似性,推测这两种植物的亲缘关系可能较近,而且萝藦科与夹竹桃科中一些植物的系统位置需进一步的研究。(4)本研究结果支持仔榄树属应从山橙族中分离出来独立成族,海杧果属应从萝芙木族中独立出来的观点;但不支持新的分类方法中将糖胶树和盆架树同归属于鸡骨常山属的观点。     

21种中国荚蒾属植物叶表皮微形态特征研究北大核心CSCD

为解决荚蒾属植物分类学问题并探究叶片解剖结构在分类学中的价值，该研究利用光学显微镜和扫描电子显微镜对21种中国荚蒾属植物的叶表皮细胞、气孔大小、气孔器类型和角质层纹饰等叶表皮微形态特征进行了观察和比较，为荚蒾属种间分类关系提供证据。结果显示，叶表皮细胞有无规则形和多边形2种，垂周壁式样有平直、浅波状、波状与深波状4种式样，其中下表皮细胞形状均为不规则形，且垂周壁式样较上表皮更稳定。气孔均分布在下表皮，形状为近圆形、椭圆形2种；气孔器类型包括不等型、平列型和无规则型3种；气孔器外拱盖内缘有近平滑、浅波状、波状3种；气孔器外围角质层纹饰有脊状隆起、条状隆起2种。研究表明，荚蒾属植物叶表皮细胞和气孔器特征在种间差异明显，可作为种间鉴定及分类的重要依据，具有重要的分类学意义。     

安徽黄精属(Polygonatum)植物叶表皮研究

利用光学显微镜及扫描电子显微镜观察了安徽省黄精属９种植物的叶表皮;统计并测量了气孔类型、气孔大小、气孔密度及气孔指数等;描述了表皮角质膜、蜡质纹饰、气孔的形状及气孔外拱盖等有关特征。结果表明：黄精属植物叶表皮气孔器类型属只有２个保卫细胞而无任何副卫细胞的单子叶植物气孔器类型。气孔的某些特征,如气孔器类型、气孔器分布特征、表皮细胞形状及垂周壁式样、角质层及蜡质纹饰等在种间差异不大,可作种间区别的次级特征用。     

苦丁茶冬青及其近缘种叶表皮微形态特征与分类学意义

利用光学显微镜和扫描电镜对冬青属苦丁茶冬青(Ilex kudingcha C.J.T seng)与5个近缘种和一形态居间物种的叶表皮微形态特征进行了观察,结果显示:(1)7种冬青属苦丁茶植物的叶表皮微形态特征具有一定的相似性,如气孔器均位于下表皮,每种植物均有正常气孔器和大气孔器,气孔器类型均以环列型为主,其次为双环列型。(2)7种冬青属苦丁茶植物的差异性体现在叶表皮细胞的垂周壁形态和气孔外缘角质层纹饰等方面。(3)霍山冬青叶表皮微形态与华中枸骨存在着较大区别。(4)结合分子生物学证据与叶表皮特征,形态居间物种类型应为五棱苦丁茶的变种。研究认为叶表皮微形态特征具有种水平上的分类学意义,能够为苦丁茶冬青及其近缘种的划分提供可靠的依据。     

新疆14种青兰属植物叶表皮微形态结构研究

应用光学显微镜和扫描电镜,对新疆14种青兰属(DracocephalumL.)植物叶片的微形态特征进行观察,结果表明:该属植物只有宽齿青兰(Dracocephalum paulsenii Briq.)叶的下表皮细胞为多边形,其余种植物叶的上、下表皮均为不规则形;垂周壁式样有深波状、波状、浅波状、平直-弓形;气孔器均为横列型;气孔形状有椭圆形、长椭圆形、宽椭圆形、近圆形;表皮毛的类型为小刺毛、短柔毛、长柔毛、头状腺毛和盾状腺毛。在不同种间甚至是同种植物叶的上、下表皮上,气孔器的大小、气孔密度、气孔指数、气孔外拱盖内缘、表皮毛、角质层纹饰等的微形态特征都存在很大的差异,可为探讨本属种间的分类、亲缘关系和演化趋势提供理论依据。     

中国蔷薇属植物叶表皮微形态特征及其系统学意义

应用光学显微镜和电子显微镜,对30种中国蔷薇属植物叶表皮的微形态特征进行了观察。结果表明:叶表皮细胞一般为多边形或不规则形,垂周壁平直、平直弓状、波状、浅波状和深波状式样;该属植物的叶表皮细胞大小种间差异较大,总体来看上表皮几乎全为多边形细胞,且上表皮细胞比下表皮细胞略大;在扫描电镜下观察,大部分细胞的平周壁下陷,仅部分细胞的平周壁突起;叶表皮角质层纹饰多样、即使同一个种的上下表皮细胞角质层纹饰微形态特征也有差异;大多植物叶片表面具柔毛,均为单毛,有长柔毛和短柔毛,大部分种类的柔毛基部无特化,其基部由普通的表皮细胞围绕着,少部分种类的柔毛基部由几个特化的辐射状细胞围绕着,少数植物无毛;有些植物叶表面或叶脉上还分布有腺毛;除了单叶蔷薇亚属的小蘗叶蔷薇1个种的上下表皮均分布有气孔器外,其余蔷薇亚属的29种植物的气孔器均分布在下表皮,形状为椭圆形、宽椭圆形和长椭圆形三种,气孔器类型有无规则型、不规则四细胞型、对位四细胞型、环列型和辐射型;气孔器外拱盖内缘近平滑或呈深浅不同的波状、气孔器外拱盖纹饰光滑或粗糙,大多植物气孔器的保卫细胞两级T型加厚;另外,只有小蘗叶蔷薇的气孔器具双层外拱盖且仅上表皮被毛等特征也说明了其在演化上的特殊地位,蔷薇属植物叶表皮的这些微形态特征,在属内各组间无明确的规律性,但可为探讨该属种间的分类学及亲缘关系提供依据。     

观音座莲目的角质层特征

运用扫描电镜和光学显微镜对观音座莲目4种(属)植物的叶角质层进行了研究。结果表明,其间存在明显共性:表皮构造均为单面气孔式,上下表皮细胞垂周壁皆呈不同程度的波状起伏,气孔器类型皆为环列型。需要指出的是观音座莲目的全部6属植物均具环列型的气孔器,这表明该特征在目一级水平上具有重要的分类学及系统学意义。同时天星蕨属的角质层特征进一步支持了将天星蕨属上升为天星蕨科的观点。     

新疆12种黄芩属植物叶表皮微形态结构的研究

应用光学显微镜和扫描电镜技术对新疆12种黄芩属植物叶片上的微形态特征进行观察。结果表明:该属植物叶的上表皮细胞形状及垂周壁式样有多种形式;而下表皮细胞形状均为不规则形,垂周壁式样均为深波形,不具分类学意义,但叶片两面分布的气孔器,在不同种间气孔大小、气孔密度、气孔指数、气孔外拱盖内缘等方面都存在着显著差异;其表皮角质层纹饰和表皮毛的微形态也各有不同;大多数植物叶片表面具腺点,其大小、分布及疏密程度也有不同。植物叶表皮上的这些微形态特征,可为探讨本属种间的分类学及亲缘关系提供一定的佐证。     

中国罗汉松属叶角质层微形态结构及其分类意义

利用扫描电镜对罗汉松属8种2变种植物叶角质层内外表面进行了细致观察。发现罗汉松属植物叶角质层结构具有许多相似特征,表皮细胞较为规则,长方形或多边形,边缘波状弯曲;气孔器排列成带状,长轴均与叶脉一致,气孔器具较为明显的气孔塞和伏罗林环,气孔器保卫细胞极延伸明显,通常具有2~4个副卫细胞、不具极副卫细胞。但罗汉松属叶角质层结构也具有明显的种间差异,镰叶罗汉松和洛杉矶罗汉松同其它种类差异最大,这两种植物叶两面均具气孔器,角质层内表面垂周壁直,角质层凸缘不明显;贺氏罗汉松最为显著的特征是近轴面和远轴面表皮细胞的垂周壁角质层厚且凸缘均极其发达;小叶罗汉松近轴面表皮细胞排列较为规则,多数为方形,长轴与叶脉垂直,垂周壁之间的角质层突起较为显著,延伸到皮下层;兰屿罗汉松近轴面表皮细胞排列较不规则,多边形,细胞的角端比较钝,没有棱角;大理罗汉松气孔带间隔较小,有时两条气孔带挤在一起,使副卫细胞紧连,近轴面表皮细胞较短,方形或长方形,垂周壁之间的角质层较不发达;海南罗汉松角质层气孔带间隔较宽,气孔器形状为阔椭圆形,近轴面表皮细胞均为细长方形;变种短叶罗汉松和狭叶罗汉松与罗汉松也具有明显差异,短叶罗汉松近轴面表皮细胞排列不规则,垂周壁深波状弯曲,凸缘极为明显,但原种罗汉松近轴面表皮细胞排列较为规则,垂周壁浅波状弯曲,凸缘不明显,而狭叶罗汉松近轴面表皮细胞方形或长方形,比罗汉松的表皮细胞短,垂周壁直或略弯曲,角质层极厚。这些角质层微形态特征差异可以作为罗汉松属内种类分类鉴定的依据。     

24种鳞毛蕨科植物叶表皮形态特征的研究

该研究利用光学显微镜对鳞毛蕨科24种植物的叶表皮形态特征进行观察。结果表明:(1)24种鳞毛蕨科植物的上表皮细胞形状为长条形或不规则形,垂周壁为深波状或浅波状,下表皮细胞均为无规则形,垂周壁均为深波状;上表皮细胞长宽比在1.5~5.7之间,下表皮细胞长宽比在2.2~3.9之间。(2)在24种鳞毛蕨科植物中共观察到8种气孔器类型,分别为不等细胞型、无规则四细胞型、极细胞型、腋下细胞型、横列型、无规则型、聚腋下细胞型和聚合极细胞型,每种植物具有2~8种气孔器类型,气孔均为下生型,多为椭圆形;气孔的长宽比在1.2~1.8之间,气孔密度在17.4~86.0个/mm~2之间,气孔指数为8.60%~37.4%。(3)通过对24种鳞毛蕨科植物的观察可将其上表皮细胞形状、垂周壁形状、上表皮细胞长宽比、主要气孔器类型及衍生类型等作为叶表皮形态特征的分类依据。(4)根据叶表皮形态特征可将24种鳞毛蕨科植物分为2类:即耳蕨类和鳞毛蕨类。该研究在一定程度上支持秦仁昌分类系统对鳞毛蕨科的划分,为鳞毛蕨科植物的系统分类及演化研究提供基础资料。     

6种松科植物叶表皮的扫描电镜观察

应用扫描电子显微镜,观察了松科(Pinaceae)3个属6种植物叶表皮特征。金钱松气孔带位下表面,气孔区与非气孔区表皮细胞形态差异很大,气孔细缝状;雪松针叶四面都具气孔线,气孔下陷,孔口光滑,针叶表面具较厚的角质层,表皮细胞形状观察不清;五针松针叶仅两腹面有气孔线,气孔有隆起的圆环围绕,表面角质层呈条纹状排列,形成一种沟槽;黑松、湿地松、马尾松针叶腹背面都有气孔线,气孔呈蜂窝状排列成行,气孔下陷,孔口角质化强烈。结果表明松科3个属植物叶表皮的形态结构及气孔器差异显著,一定程度上证实了3属是自然的分类群。     

PENETRATION OF 2,4-D IN RELATION TO CUTICLE THICKNESS

Cuticle thickness was measured, either by direct microscopic examination or as weight per unit area, for the astomatous cuticle from the upper leaf surface of nine species and from tomato fruit. Thickness ranged from 1.4 μm for peach leaf cuticle to 10.8 μm for oleander leaf cuticle, and the weight from 0.19 mg/cm² to 1.26 mg/cm², respectively. Cuticles were isolated by the pectinase method and permeability to 2,4-D was determined. There was no correlation between cuticle thickness and penetration of 2,4-D, either for non-dewaxed cuticles or after chloroform extraction of waxes. Penetration of 2,4-D was increased following wax removal, but there was no correlation between wax content and the magnitude of the increase. It is suggested that cutin and wax qualitative composition are probably more important than thickness in determining relative permeability of cuticle from different plant species to 2,4-D.     

Dispersed cuticles from the Eocene of North America

KOVACH, W. L. & DILCHER, D. L., 1984. Dispersed cuticles from the Eocene of North America. Macerations of organic-rich clay from the Claiborne Formation (Middle Eocene) of Tennessee have yielded a wide variety of well-preserved dispersed cuticles. Details of the epidermal cell patterns, arrangement of the stomata1 complex, trichomes and trichome attachment have made possible the association of some dispersed cuticles with leaf types known from this formation which have similar cuticle, and with modern families. All dispersed cuticles are classified in a morphologic system. Through both our own work and a review of previous investigations we have found that distinct dispersed cuticle types can be recognized and may be used biostratigraphically to characterize geologic strata, palaeo-ecologically to provide insights into environmental reconstruction, and systematically to follow the history of certain taxa.     

The presence of cutan limits the interpretation of cuticular chemistry and structure: Ficus elastica leaf as an example

Plant cuticles have been traditionally classified on the basis of their ultrastructure, with certain chemical composition assumptions. However, the nature of the plant cuticle may be misinterpreted in the prevailing model, which was established more than 150 years ago. Using the adaxial leaf cuticle of Ficus elastica, a study was conducted with the aim of analyzing cuticular ultrastructure, chemical composition and the potential relationship between structure and chemistry. Gradual chemical extractions and diverse analytical and microscopic techniques were performed on isolated leaf cuticles of two different stages of development (i.e. young and mature leaves). Evidence for the presence of cutan in F. elastica leaf cuticles has been gained after chemical treatments and tissue analysis by infrared spectroscopy and electron microscopy. Significant calcium, boron and silicon concentrations were also measured in the cuticle of this species. Such mineral elements which are often found in plant cell walls may play a structural role and their presence in isolated cuticles further supports the interpretation of the cuticle as the most external region of the epidermal cell wall. The complex and heterogeneous nature of the cuticle, and constraints associated with current analytical procedures may limit the chance for establishing a relationship between cuticle chemical composition and structure also in relation to organ ontogeny.     

分散角质层的分类和命名*

简略讨论了分散角质层对地层学、古生物学和古环境的意义;简述了气孔器的形态学、发生学和形态一发生学的分类;在介绍并讨论了分散角质层的分类原则和命名方法的基础上,提出了新的分类方案.     

Penetration of chemicals into the Malus leaf cuticle

The adaxial leaf cuticle of Malus pumila was examined by electron microscopy to determine possible avenues for transcuticular movement of foliarly applied chemicals. Cutin-embedded polysaccharide microfibrils originated at the outer epidermal cell wall and occasionally extended to the cuticle surface. Lamellae, ca. 4 nm wide, usually were oriented parallel to the cuticle surface. When oriented perpendicular to the surface, they extended nearly to the subjacent wall layer from the surface. Aqueous solutions of uranyl acetate, silver nitrate and phenyl mercuric acetate applied to the cuticle surface of leaf segments floated on solutions of phosphate salts or thiocarbohydrazide (TCH) reacted within the cuticle to form insoluble electron-opaque deposits indicative of their avenues of transcuticular movement. Uranyl phosphate deposits were observed only in the polysaccharide microfibrils of chloroform: methanolextracted leaves. Silver-TCH deposits were observed in the microfibrils of both extracted and nonextracted leaf cuticles. Phenyl mercuric acetate-TCH deposits were randomly dispersed throughout the extracted cuticle and not associated with the polysaccharide microfibrils.Abbreviations TCH thiocarbohydrazide - PMA phenyl mercuric acetate     

Investigations on epiphytic living Pseudomonas species from Malus domestica with an antagonistic effect to Venturia inaequalis on isolated plant cuticle membranes

In order to understand better the survival and mutual interaction of epiphytic bacteria and fungi on apple plants, bacteria collected from these plants were cultivated on intact adaxial, stoma free cuticle membranes originally obtained from apple. The bacteria were labelled with luciferase genes from Vibrio harveyi in order to follow up their development and activity on the isolated cuticles. Our finding was that the epiphytic bacteria can have access to nutrients below the cuticle without causing damage to these cuticular membranes. Bacterial proteins may enable this nutrient mobilization and we found, indeed, that more than 46 proteins that must have been delivered by the bacteria in response to interaction with the cuticles as they could be found below the cuticle membrane. Eight major representatives of this group of external proteins have been sequenced with electron spray quadrupole time of flight mass spectrometry and subsequently identified by data base homology search as a flagellin, a porin type protein and proteins that are involved in amino acid recruitment and metabolism.     

The importance of leaf cuticle for carbon economy and mechanical strength

Cuticle thickness of leaves varies >?100 times across species, yet its dry mass cost and ecological benefits are poorly understood. It has been repeatedly demonstrated that thicker cuticle is not superior as a water barrier, implying that other functions must be important. Here, we measured the mechanical properties, dry mass and density of isolated cuticle from 13 evergreen woody species of Australian forests. Summed adaxial and abaxial cuticle membrane mass per unit leaf area (CMA) varied from 2.95 to 27.4?g m(-2) across species, and accounted for 6.7-24% of lamina dry mass. Density of cuticle varied only from 1.04 to 1.24?g?cm(-3) ; thus variation in CMA was mostly due to variation in cuticle thickness. Thicker cuticle was more resistant to tearing. Tensile strength and modulus of elasticity of cuticle were much higher than those of leaf laminas, with significant differences between adaxial and abaxial cuticles. While cuticle membranes were thin, they could account for a significant fraction of leaf dry mass due to their high density. The substantial cost of thicker cuticle is probably offset by increased mechanical resistance which might confer longer leaf lifespans among evergreen species.     

Patterns of Effective Permeability of Leaf Cuticles to Acids

Plants in the field are frequently exposed to anthropogenic acid precipitation with pH values of 4 and below. For the acid to directly affect leaf tissues, it must pass through the leaf cuticle, but little is known about the permeability of cuticles to protons, or about the effect of different anions on this permeability. We investigated the movement of protons through isolated astomatous leaf cuticles of grapefruit (Citrus X paradisi Macfady.), rough lemon (Citrus limon [L.] Burm. fils cv Ponderosa), and pear (Pyrus communis L.) using hydrochloric, sulfuric, and nitric acids. Cuticles were enzymically isolated from leaves and placed in a diffusion apparatus with pH 4 acid on the morphological outer surface of the cuticle and degassed distilled water on the inner surface. Changes in pH of the solution on the inner surface were used to determine rates of effective permeability of the cuticles to the protons of these acids. Most cuticles exhibited an initial low permeability, lasting hours to days, then after a short transition displayed a significantly higher permeability, which persisted until equilibrium was approached. The change in effective permeability appears to be reversible. Effective permeabilities were higher for sulfuric acid than for the others. A model of the movement of protons through the cuticle is presented, proposing that dissociated acid groups in channels within the cutin are first protonated by the acid, accounting for the low initial effective permeability; then protons pass freely through the channels, resulting in a higher effective permeability.     

新疆亚鳞木（比较种）角质层特征

报道了产自江苏宜兴晚泥盆世五通组新疆亚鳞木（比较种）Ｓｕｂｌｅｐｉｄｏｄｅｎｄｒｏｎ ｃｆ． ｘｉｎｊｉａｎｇｅｎｓｅＳｕｎ）的表皮角质层用荧光分析显示的特征．该种茎干表皮角质层覆于叶座及叶座间隔带,其中间隔带角质层厚于叶座．表皮细胞在间隔带与叶座表现特征不同．间隔带中部,表皮细胞呈纵长的多边形,其纵长方向与茎干延伸方向相同,细胞壁略有弯曲．间隔带靠近叶座之表皮细胞,细胞壁直,形状类似于前者;但大小仅为前者的１／２左右,且其纵长方向逐渐向叶座边缘偏转．叶座表皮细胞呈近等多边形,有胞间隙．该种茎表皮无气孔