Variation in the structure and development of foliar stomata in the Euphorbiaceae

The structure and ontogeny of foliar stomata were studied in 50 species of 28 genera belonging to 17 tribes of the family Euphorbiaceae. The epidermal cells are either polygonal, trapezoidal, or variously elongated in different directions and diffusely arranged. The epidermal anticlinal walls are either straight, arched or sinuous. The architecture of cuticular striations varies with species. The mature stomata are paracytic (most common), anisocytic, anomocytic and diacytic. Occasionally a stoma may be tetracytic, cyclocytic or with a single subsidiary cell. The ontogeny of paracytic stomata is mesogenous dolabrate or trilabrate, mesoperigenous dolabrate; that of diacytic stomata is mesogenous dolabrate, whereas that of anisocytic stomata is mesogenous trilabrate; rarely an anisocytic stoma may be mesoperigenous. Hemiparacytic stomata are mesoperigenous unilabrate; tetracytic stomata are mesoperigenous dolabrate and anomocytic stomata perigenous. Abnormalities encountered include four types of contiguous stomata, stomata with a single or both guard cells aborted and persistent stomatal initials. Cytoplasmic connections between the guard cells of two adjacent stomata or the guard cell of a stoma and an adjacent epidermal/subsidiary cell, or both types occurring in a species, were noticed. The stomatal development, distribution, diversity and basic stomatal type with reference to systematics are discussed.     

27种木犀属植物叶表皮微形态特征的研究

利用光学显微镜和扫描电子显微镜观察了27种木犀属植物的叶表皮,测量并统计了气孔器类型、气孔大小、气孔密度、气孔指数及腺点密度等指标,以明确各种的叶表皮细胞形状及垂周壁式样、表皮角质膜、蜡质纹饰及气孔外拱盖的具体特征。结果显示:木犀属植物叶下表皮有气孔器,形状为圆形、椭圆形;气孔器类型多为不规则型,只有总状桂花和狭叶木犀为环列型;气孔器外围角质层有放射状、条状、环状、颗粒状等多种类型;叶表皮细胞形状有无规则形和多边形2种;下表皮腺点的数目远远大于上表皮。研究表明,木犀属植物气孔器和叶表皮细胞特征在种之间差异比较明显,可以作为种间鉴定的重要依据,具有重要的分类学意义。     

Epidermal studies in some Indian cultivars of Bougainvilleas

Epidermal studies in fifteen Indian cultivars of Bougainvilleas are described. The epidermal cells are polygonal isodiametric, or elongated with thick straight arched or slightly sinuous walls. Parallel culticular striations are radiating from guard cells. The mature stomata are anomocytic, paracytic and with a single subsidiary cell. The abnormal types noticed are: single guard cells with or without pores, arrested development, variously oriented contiguous stomata, cytoplasmic connections between nearby stomata and epidermal or subsidiary cells, and persistent stomatal cells. The development of anomocytic stomata is perigenous while that of the other types is mesogenous. Fifteen cultivars of Bougainvilleas are separated on the basis of bract colour, stomatal frequency and index per unit area.     

天南星科叶表皮研究

利用光学显微镜对天南星科18属27种及菖蒲科1属1种植物的叶表皮微形态进行观察,同时用扫描电镜对具代表性的14种植物作了研究,结果显示：天南星科气孔类型变异较大,有不规则型,辐射型,平列型,胞环型及平列型和胞环型间的过渡类型,副卫细胞数目0－12个;表皮细胞长宽近相等,平周壁具条纹或否,垂周壁平直,弧形或波浪形,虽然气孔类型对天南星科分类上的意义不大,但与表皮细胞垂周壁形状,副卫细胞角质层纹饰等特征相结合对种间分类有一定意义,天南星科与菖蒲科叶表皮微形态明显不同,从而支持菖蒲属从天南星科中分出另立为科的观点。     

Morphological characters of leaf epidermis in schisandraceae and their systematic significance

The leaf epidermis of 23 species belonging to 2 genera within Schisandraceae was investigated using light and scanning electron microscopy. Many characters of the leaf epidermis in Schisandraceae, such as shape of epidermal cells, type of stomata, and cuticular ornamentation, are usually constant within species and thus helpful for elucidating the relationship between and within genera. Leaf epidermal cells are usually irregular or polygonal in shape. The patterns of anticlinal walls are straight, sinuolate, sinuous or sinuate. The stomatal apparatus belong to paracytic or laterocytic type and the latter is subdivided into various subtypes based on the number and arrangement of subsidiary cells. Under scanning electron microscopy observation, the cuticular membrane is often striated, sometimes squamulate or granular; the inner margin of the outer stomatal rim is nearly smooth or denticulate. Evidences from shape of epidermal cells, patterns of cuticular intrusions between the ends of each guard cell of a pair and distribution of stomatal apparatuses support the viewpoint thatKadsura is more primitive thanSchisandra. Study on leaf epidermis also shows thatKadsura interior deserves the rank of a distinct species and the treatment of the evergreen groups, includingS. propinqua andS. plena, as distinct from the deciduous species of the genus is quite natural.     

中国桑寄生科植物叶表皮微形态

通过扫描电镜对中国桑寄生科桑寄生亚科8属18种和槲寄生亚科1属2种植物成熟叶的上、下叶表皮内表面和下表皮外表面进行了研究。内面观发现桑寄生科植物叶上、下表皮形状为多边形,垂周壁式样平直或稍弓形,常具有角质增厚,平周壁常覆盖厚角质或颗粒状、丝状角质增厚;气孔存在于上下表皮,通常下表皮较多,气孔的形状,特别是保卫细胞的形态在亚科间、属间或种间都具有一定的差异,气孔器类型为平列型或单圈型。下表皮表面观察了的角质膜和蜡质纹饰、气孔的形状,外部气孔缘及外部气孔缘内缘的特征。这些特征在亚科或属级水平上较为稳定,有的也表现出种间差异,有一定的分类价值。从气孔形态和外部气孔周围角质膜来看,两亚科显示出明显的不同:桑寄生亚科上、下表皮均具有内部气孔缘,而槲寄生亚科没有此结构;桑寄生亚科外部气孔周围角质膜增厚成环状,其上具增厚的条纹,而槲寄生亚科外部气孔周围角质膜增厚成脊状,不具条纹。这些特征支持槲寄生亚科作为独立1个科来处理。     

Cuticle micromorphology of leaves of Pinus (Pinaceae) from Mexico and Central America

Cuticle micromorphology of 34 taxa of Pinus from Mexico and Central America was studied with scanning electron microscopy, and leaf morphology was described. In total, 29 characters, 22 from the inner cuticular surfaces and seven from the outer, were described in detail. These characters have value either for testing infragenerie classifications or for identifying individual taxa. Characters relating to the periclinal wall texture of the epidermal cells, the shape and degree of development of the anticlinal walls of the epidermal cells, the basal and apical shapes of anticlinal epidermal cell walls, the continuity of the epidermal cells, the size ratio of the polar to lateral subsidiary cells, the grooves on subsidiary cells, the cuticular flanges between guard and subsidiary cells, the groove near the bristles and the elevation of the Florin ring ridge and striations on the Florin ring are particularly useful for infrageneric classification. The agreement between these characters and infrageneric classifications is discussed. Characters relating to the end wall shapes of the epidermal cells, the relative length of epidermal cells, the shape of the stomatal apparatus, the texture of guard and lateral subsidiary cell surfaces, the polar extensions, the number of subsidiary cells and epidermal cell layers between stomatal rows, the integrity of stomatal rows, cell numbers between stomata in a row, cuticular flanges between guard cells, bristle flanges and surface textures, epicuticular waxes, striations on Florin rings and stomatal shapes, contain some important information for identifying Mexican pines. The distribution of the states of each character is compared with that of the Asian pines. Cuticular characters are used to help determine the affinities of taxonomically difficult taxa.     

The stomatal apparatus of Lycopodium japonicum and its bearing on the stomata of the Devonian lycophyte Drepanophycus spinaeformis

The structure of the stomatal apparatus of the leaf of Lycopodium japonicum Thumb was studied using epidermal macerations, sections and scanning electron microscopy. The stomatal apparatus of L. japonicum consists of two large guard cells and pore, and is anomocytic. Based on light microscopy, the impression from epidermal macerations that there were two small guard cells surrounded by two, large, similarly shaped, subsidiary cells (paracytic) derives from a pronounced elliptical cuticular ledge on the surface of the guard cells surrounding a thickened circumporal area. A similar appearance is characteristic of cuticle preparations of the Devonian lycophyte Drepanophycus spinaeformis Göppert. We therefore conclude, as did W.H. Lang over 70 years ago, that the stomata of the early lycophyte were also anomocytic, as were those of a second species of Drepanophycus , D. qujingensis Li & Edwards.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 209–216.     

Cuticle micromorphology of leaves of Pinus (Pinaceae) in east and south-east Asia

Cuticle micromorphology of 21 species of Pinus in east and south-east Asia was studied with scanning electron microscopy and leaf morphology was described. Cuticle characters that can be distinguished by their size, shape, and surface texture, as well as by the number of subsidiary cells, the rows of epidermal cells within a stomatal band, the number of cells between stomata in stomatal rows, and the patterns of cuticular flanges are described in detail. Most of these characters have not been considered in sufficient detail previously. Some of the inner cuticular features, such as the texture of periclinal walls, the shape of the top of the anticlinal walls, the comparative size of the lateral and polar subsidiary cells, and the development of the groove near bristles, are diagnostic characters for subgenera. The agreement between cuticle characters and taxa is discussed. Cuticular characters were used to try to resolve the position of taxa that currently appear in different places in different classifications.     

中国梅花草属植物的叶表皮特征及其系统学意义

利用光学显微镜和扫描电镜对梅花草属Parnassia 30种植物的叶表皮进行了观察。结果表明:气孔器普遍存在于叶的下表皮,少数种的上表皮也有分布,均为无规则型。叶表皮细胞形状为多边形或不规则形;垂周壁式样可区分为近平直、浅波状和波状。在扫描电镜下,叶表皮气孔器外拱盖内缘为近平滑、浅波状或波状;一些种的保卫细胞两端有加厚;角质膜条纹状,有的条纹隆起,有的条纹上附有颗粒或小孔穴。气孔器类型及下表皮细胞形状的一致性表明梅花草属是一个自然分类群;sect. Saxifragastrum叶表皮特征具有多样性显示该组可能是一个复合群;突隔梅花草P. delavayi属于subsect. Xiphosandra,其气孔下陷,与其细胞学特征相似,支持独立为一组;此外,气孔器的分布、保卫细胞两端加厚、气孔器外拱盖内缘形态以及角质膜等特征对该属部分种的区分有一定的参考价值。     

Epidermal Structure and Ontogeny of Stomata in some Verbenaceae

The present paper deals with epidermal structure and developmentof stomata in 14 species of Verbenaceae. The epidermal cellsare either polygonal, isodia-metric, or elongated in variousdirections, and irregularly arranged. The anticlinal walls arethick, mostly sinuous, occasionally arched or straight. Thesurface of the cuticle shows parallel, rarely corrugated, striations.Some 12 types of eglandular and glandular trichomes, and foliarnectaries are noticed. The mature stomata are diacytic, anisocytic,paracytic, with a single subsidiary cell, anomocytic and perigenous.The development of anomocytic stomata is perigenous, while thatof others is mesogenous or syndetocheilic type. Abnormalitiesnoticed here include contiguous stomata, stomata with a singleguard cell, and aborted guard cells.     

Leaf epidermal morphology of Zanthoxylum s.l. (Rutaceae) from China

Leaf epidermis characters in 40 of ca. 250 species of Zanthoxylum (Rutaceae) were investigated using light microscopy (LM) and scanning electron microscopy (SEM). The stomata are anomocytic and exist only on the abaxial epidermis except in Z. nitidum, which also has stomata on the adaxial surface. The epidermal cells are usually polygonal or irregular in shape, with anticlinal walls straight, arched, repand, or sinuous. Under the SEM, the inner margin of the outer stomatal rim is nearly smooth, sinuolate or erose, and the cuticular membrane of the leaf epidermis is smooth, striate, or sometimes striate to wrinkled. These data of leaf epidermis of Zanthoxylum demonstrated that there exist many common characters between subgen. Fagara and subgen. Zanthoxylum, sug-gesting a close relationship between the two subgenera. The utility of some characters in identifying some species of Zanthoxylum was also discussed.     

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。     

栒子属（Cotoneaster Medikus）植物叶表皮微形态特征及其分类学意义

在光学显微镜和扫描电镜下,对栒子属（Cotoneaster Medikus）2组15种植物的叶表皮特征进行了观察,发现疏花组（Sect. Cotoneaster）植物叶表皮细胞多为多边形,垂周壁一般为平直—弓形,气孔均为无规则型;单花组（Sect. Uniflos）植物叶表皮细胞常不规则型,垂周壁多浅波状,气孔除无规则型（anomocytic）外还兼有十字型（staurocytic）、四分体型（tetracytic）和等三体型（isotricytic）。依据气孔周围角质膜特征等叶表皮性状,把15种植物分为W型、S型和I型,其中W型是S型的特化类型,疏花组的种类为W型或S型,单花组的种类为I型。叶表皮微形态特征可以为栒子属植物组及种的分类学处理提供解剖学资料。     

A systematic investigation of leaf epidermis in <Emphasis Type="Italic">Camellia</Emphasis> using light microscopy

The characters of mature leaf epidermis of 58 species and 3 varieties belonging to 19 sections, representing all the 4 subgenera: Protocamellia, Camellia, Thea and Metacamellia were investigated under light microscope. The main conclusions are as follows: 1) The shape of the epidermal cells is elliptical, polygonal or irregular and the anticlinal walls are arched, straight or sinuolate, sinuous to sinuate. 2) The stomatal apparatus, consisting of a stoma (a pore plus a pair of guard cells) and 3–4 subsidiary cells, restricted to the abaxial surface of the leaves, were assigned to anisocytic, anisotricytic, isotricytic or tetracytic type. 3) Special structures in leaf epidermis, such as stomatal clusters, aborted stomata, secretory cells, cuticular intrusions and their taxonomic implications were also elucidated.     

Observations on the cotyledonary and hypocotyledonary stomata and trichomes in some Caesalpiniaceae with a note on their taxonomic

The structure and ontogeny of stomata on cotyledons and hypocotyls and the trichomes on hypocotyl are accounted for in eighteen species of Caesalpiniaceae. Trichomes are eglandular, bi- rarely tri-celled, smooth walled or walls wavy with cuticular striations or tubercles. Anomocytic, haplocytic, paracytic, diacytic, transitional, tetracytic, tricytic and cyclocytic stomata occur in different combinations even on the same surface of the cotyledon. In all, there are fourteen combinations. Inspite of the diversity, the most frequent type is anomocytic in most of the species and paracytic in some species of Cassia and Delonix (abaxial) but rarely it is haplocytic or anisocytic. In hypocotyls it is anomocytic. Ontogenetically anomocytic, tetracytic and cyclocytic stomata are perigenous, whereas other types are mesogenous or mesoperigenous. There is an increase in the number of subsidiary cells by their division or the neighbouring perigenes assuming their shapes. About eight such types are described. A pair of stomata has a common subsidiary cell. Twelve types of guard cell and stomatal approximation abnormalities are described. A range in the number, size and shape of the nuclei in guard cell are recorded. Megastomata (giant stomata) are observed in Parkinsonia and Tamarindus. The taxonomic significance of the stomata is also discussed.     

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。     

安徽景天属（Sedum）植物叶表皮研究

利用光学显微镜和扫描电子显微镜观察安徽省１２种景天属（Ｓｅｄｕｍ）植物的叶表皮,统计并测量报气孔类型,气孔大小,气孔密度及气孔指数等,描述了表皮角质膜,蜡质纹饰及气孔外拱盖的有关特征。结果表明：景天属植物叶表皮气孔器为不等细胞型。气孔器类型,表皮细胞形状及垂周壁式样,角质膜,蜡质纹饰等,种间无差异或极小,而气孔大小,气孔器分布特征,气孔密度,表皮毛等种间差异较大,建议在种间区分上将差异大的特征作为     

Epidermal Structure and Ontogeny of Stomata in Vegetative and Reproductive Organs of Ephedra and Gnetum

The epidermal structure and development of stomata in vegetativeand reproductive organs of Ephedra foliata and Gnetum ula isdescribed. The epidermal cells are polygonal, isodiametric,or elongated with thick or thin straight, arched, or slightlysinuous anticlinal walls. The cuticle is thin or thick. Papilla-likeunicellular outgrowths are present in Ephedra foliata. The maturestomata are orientated parallel to the longitudinal axes orirregularly. The mature stomata are anomocytic, paracytic, witha single subsidiary cell, cyclocytic, and actinocytic. Arresteddevelopment, contiguous stomata, and stomata with aborted guardcells have been observed. The ontogeny of stomata on differentorgans of these two plants is typically haplocheilic or perigenousbut the stomatal apparatus varies from organ to organ.     

The stomata of the cork-oak, Quercus suber. An ultrastructural approach

In the evergreen leaves of Quercus suber, stomata play a major role in adaptation to drought and temperature stress. The leaf is of zygostomic type and has about 430 stomata per square milimeter of abaxial leaf surface. The stomatal complex is of the anomocytic type. The guard cells protrude from the epidermal plane. The guard cell nucleus contains heterochromatin in small granules. The guard cell cytoplasm is characterised by a large number of well developed mitochondria, amyloplasts with stroma and grana, and a well developed cytoskeleton with a cortical array of microtubules oriented pa railed to the slit axis that persist even in mature cells. Guard cell walls are asymmetrically thickened and devoid of plasmodesmata. No area of cell walls was free of cuticle or covered by a thin cuticular layer and apparently no area of limited cuticular development provides evaporation when the stomata are closed.