Structure and function of trichornes in the shoot tip of Ardisia crispa (Thunb.) A.DC. (Myrsinaceae)

MILLER, I. M., GARDNER, I. C. & SCOTT, A., 1984. Structure and function of trichomes in the shoot tip of Ardisia crispa (Thunb.) A.DC. (Myrsinaceae). The trichomes in the shoot tip of the myrsinaceous leaf-nodulated species Ardisia crispa (Thunb.) A.DC. have been studied at the ultrastructural level. Two distinct kinds of trichomes are found. Small peltate scales arise from the abaxial protoderrn of the developing leaves. Multicellular stellate trichomes on the adaxial surface of young leaves are uniseriate, rotate and consist of up to eight multicellular arms radiating from a central stalk. The multicellular arms terminate distally in swollen club-shaped tips. The stellate trichomes secrete large quantities of carbohydrate-containing mucilage. Colonies of leaf nodule bacteria are found resident in this mucilage. The role of the peltate scales and the stellate trichomes n the maintenance of the cyclic leaf nodule symbiosis is discussed.     

地黄叶和茎的解剖学及组织化学研究

采用解剖学和组织化学方法对地黄叶和茎的显微结构以及梓醇、多糖的分布进行观察研究,以明确梓醇和多糖在地黄叶和茎中的分布特征。结果显示:(1)地黄叶的上、下表皮均分布有腺毛和非腺毛,腺毛都属于头状腺毛,包括长柄和短柄的头状腺毛,两类腺毛的分泌物化学成分主要是黄酮和多糖;叶的上、下表皮上都分布有无规则型气孔,下表皮的气孔密度比上表皮的大,但气孔指数相差不大;栅栏组织由2～3层薄壁细胞构成,排列紧密,海绵组织薄壁细胞形状无规则,细胞间隙大。(2)组织化学研究表明,海绵组织中黄斑样的薄壁细胞是梓醇和多糖的贮存场所,这类薄壁细胞在叶片边缘的齿末端处最为集中,茎的皮层、韧皮部和木质部的薄壁细胞也都是梓醇和多糖的贮存场所。     

冬凌草腺毛的形态学及组织化学研究

利用光学显微镜对药用植物冬凌草地上部分腺毛的形态、分布和组织化学进行了研究。结果表明:(1)冬凌草的叶表皮有3种形态显著不同的毛,即非腺毛、盾状腺毛和头状腺毛;盾状腺毛和头状腺毛均具1个基细胞、1个柄细胞和头部;成熟的盾状腺毛的头部一般由4个分泌细胞组成,而头状腺毛头部由2个分泌细胞组成。(2)组织化学鉴定结果显示:2种腺毛中均含有黄酮类成分,盾状腺毛中还含有单萜、倍半萜等萜类成分;冬凌草甲素可能只存在于盾状腺毛中,但需要更直接的证据证明。研究认为,高密度的盾状腺毛可以作为筛选冬凌草高甲素含量品种的一项重要依据。     

Ficus carica L. leaf anatomy: Trichomes and solid inclusions

Ficus carica L., a typical plant of the Mediterranean environment, presents leaves covered by an extensive indumentum, and a mesophyll full of solid inclusions. The morphology and ultrastructure of the trichomes, calcium carbonate cystoliths and silicified structures of leaves of F. carica cv Dottato were investigated with light, confocal, scanning electron microscopy, and energy-dispersive X-ray spectroscopy. At the same time, histochemical reactions were also employed to analyse the indumentum composed by glandular and non-glandular trichomes by applying chemical reagents and fluorescence microscopy. Non-glandular and glandular trichomes, capitate, are described. Non-glandular trichomes are unicellular simple, spine-like and present different morphology and sizes. The capitate glandular trichomes are present on leaf adaxial and abaxial surface and consist of one-celled stalk and 3/4 cells spherical head. Histochemical characterisation of leaf hairs revealed the presence of flavonoids, while glandular trichome head cells showed a complex mixture of alkaloids, essential oil and flavonoids. Cu and Al were found in the constitutive structures, spike and dome, of the cystoliths. Several epidermal cells and non-glandular trichomes were silicified. Leaf hairs, trichomes secretions, solid inclusions and silicification of F. carica leaf have significant roles to play in relation to leaf protection from external factors, including high-intensity radiation, herbivores or pathogens.     

The foliar trichomes of Hypoestes aristata (Vahl) Sol. ex Roem. & Schult var aristata (Acanthaceae) a widespread medicinal plant species in tropical sub-Saharan Africa: with comments on its possible phylogenetic significance

The micromorphology of foliar trichomes of Hypoestes aristata var. aristata was studied using stereo, light and scanning microscopy (SEM). This genus belongs to the advanced angiosperm family Acanthaceae, for which few micromorphological leaf studies exist. Results revealed both glandular and non-glandular trichomes, the latter being more abundant on leaf veins, particularly on the abaxial surface of very young leaves. With leaf maturity, the density of non-glandular trichomes decreased. Glandular trichomes were rare and of two types: long-stalked capitate and globose-like peltate trichomes. Capitate trichomes were observed only on the abaxial leaf surface, while peltate trichomes were distributed on both adaxial and abaxial leaf surfaces.     

Characterization and density of trichomes on three common bean cultivars

Trichomes have been implicated as a mechanism which can confer resistance to both plant pests and drought. A study was conducted to provide information regarding genetic variability for trichome distribution and density among three diverse dry bean (Phaseolus vulgaris L.) cultivars, and to characterize the types of trichomes present among the cultivars. Trichomes on the leaf surfaces were micrographed with a scanning electron microscope (SEM) and counted using a stereomicroscope on both the abaxial and adaxial leaf surfaces of the cultivars ‘Bill Z’, ‘Pompadour Checa’ and ‘Diacol Calima’. Straight, hooked, and glandular trichomes were observed on the leaf surfaces of each cultivar. SEM micrographs are presented for the leaf surfaces of each cultivar and trichome type. The abaxial leaf surface had more straight trichomes than the adaxial leaf surface for ‘Pompadour Checa’ and ‘Diacol Calima’, however ‘Bill Z’ had more on the adaxial surface. The opposite relationship existed among the cultivars and leaf surfaces for the hooked trichomes.     

Exogenous Methyl Jasmonate Alters Trichome Density on Leaf Surfaces of Rhodes Grass (Chloris gayana Kunth)

Jasmonates, including jasmonic acid and its derivatives such as methyl jasmonate (MeJA), are plant growth substances that control various responses. Jasmonates regulate leaf trichome density in dicotyledonous plants, but their effects on the trichome density of monocotyledonous plants, such as those in the Poaceae, remain unclear. In the present study we examined the effects of exogenous MeJA on the trichome density of Rhodes grass, which has three kinds of trichomes: macrohairs, salt glands, and prickles. Exogenous MeJA significantly increased the densities of macrohairs and salt glands on the adaxial and abaxial leaf surfaces and those of prickles on the adaxial leaf surface. Because exogenous MeJA significantly reduced the leaf area, we calculated the number of trichomes per 1000 epidermal cells to eliminate the effects of reduced leaf area. Exogenous MeJA significantly increased the number of macrohairs per 1000 epidermal cells on both adaxial and abaxial leaf surfaces, but it significantly decreased the number of salt glands per 1000 epidermal cells on both surfaces. Exogenous MeJA had no significant effects on the number of prickles per 1000 epidermal cells on either of the leaf surfaces. These results indicate that exogenous MeJA alters the trichome density by affecting leaf area and trichome initiation, and the effects of exogenous MeJA on trichome initiation differ among the various trichome types.     

Micromorphology of the leaf epidermis in taxa of the Agropyron-Elymus complex (Poaceae)

WEBB, M. E. & ALMEIDA, M. T., 1990. Micromorphology of the leaf epidermis in taxa of the Agropyron-Elymus complex (Poaceae). A comparative analysis by scanning electron microscopy was carried out on both leaf epidermes (adaxial and abaxial) of Elymus pychnanthus (Godr.) Meld, and Agropyron glaucum Roemer & Schultes.
The adaxial epidermes of E.pychnanthus and A. glaucum are similar in the position and the shape of the long cells, silica bodies and costal and papillate prickles. They differ in the higher number of silica cells in E. pychnanthus and in the presence of intercostal hooks in A. glaucum.
The abaxial epidermes of both species are similar in the shape of interstomatal cells, silica bodies and papillate prickles. They differ in the distribution of the stomata and in the contact zones of the long cells. Elymus pychnanthus and A. glaucum differ also in the trichomes situated along the margins of the leaf blade.
These micromorphological differences, especially those of the abaxial epidermis, are useful taxonomic features.     

喜树各器官解剖学及其喜树碱含量测定的研究

运用植物解剖学方法研究了喜树各器官的结构,对喜树不同器官中喜树碱含量进行了HPLC法测定。结果表明:(1)喜树根尖纵切面的结构由根冠、分生区、伸长区和根毛区4个部分组成。根和茎的初生结构都由表皮、皮层和维管柱3部分组成;在根和茎初生结构的皮层和髓部具有被锇酸染成黑色的嗜锇细胞;在次生生长中,根的木栓形成层起源于中柱鞘,茎的木栓形成层起源于皮层细胞;随着次生维管组织的增加,茎的中央形成髓腔。(2)叶片由表皮、叶肉和叶脉组成,叶肉细胞中分布有嗜锇细胞;光学显微镜和扫描电子显微镜观察显示,喜树叶的上表皮没有气孔器分布;下表皮气孔器的保卫细胞呈肾形,没有副卫细胞,被几个表皮细胞不规则的围绕,属于无规则型;在上下表皮均分布着单细胞非腺毛和2种类型的腺毛,其中下表皮的分布相对稠密;上表皮腺毛呈球形,下表皮腺毛大部分为长卵形,其中间杂着一些球形腺毛;观察发现,越幼嫩的叶中,分布的腺毛和非腺毛越多。(3)喜树不同器官中喜树碱含量以幼叶和种子中较高,木质部中较低,髓中含量最少;随着叶的发育成熟,叶中喜树碱的含量逐渐降低,且同一叶的不同部位,喜树碱含量也有差异;喜树幼苗发育过程中喜树碱含量也在不断变化。     

甘草腺毛的形态发生和组织化学研究

利用扫描电镜及薄切片技术对甘草的腺毛形态发生和发育过程进行了观察,并对腺毛发育过程中黄酮类成分积累进行了组织化学定位研究。结果表明：甘草腺毛为多细胞构成的盾状腺毛,有长柄和短柄2种类型;前者主要分布在花萼片上,而后者主要分布于叶片上。组化鉴定结果显示：腺毛中存在着黄酮类成分、其他亲脂类和非纤维素多糖类成分;在腺毛的发育过程中,黄酮类物质是随腺毛的发育成熟,在头部盘状结构的分泌细胞及角质层下腔中积累。研究结果对进一步探讨甘草叶中黄酮类成分的合成及其作用提供科学依据。     

Leaf epidermis of Quercus subgen. Cyclobalanopsis (Oerst.) Schneid. (Fagaceae)

The characters of leaf epidermis (mainly epidermal cells, stomata, trichomes) have been examined in 48 species of Quercus subgen. Cyclobalanopsis of the Fagaceae. The adaxial leaf epidermis was investigated under light microscope. The epidermal cells of this surface are of two types in terms of the outline of their anticlinal wall. One type is characterized by having straight wall, and this type has been observed in most of the species studied, while the other is characterized by having sinuate or sinuolate wall, and this type has been found only in eight species, such as Q. blakei and Q. hui. There exist two types of trichome-bases on this surface: in one type the bases are singlecelled, and this type has been found in most of the species studied, such as Q. oxyodon and Q. augustinii; while in the other the bases consist of a group of cells, and this type has been found only in nine species, such as Q. thorelii and Q. chungii. The abaxial leaf epidermis was investigated under light microscope and scanning electron microscope. The epidermal cells and the trichome-bases on this surface under LM are in morphology essentially similar to those on the adaxial surface. The stomata under LM include three types: cyclocytic, subcyclocytic and anomocytic. The stomatal density is of some diagnostic value to distinguish between closely related species, e.g. the stomatal density in Q. glauca is higher than that in Q. schottkyana. Under SEM the stomata are elliptical or roundish. The most noteworthy feature of the adaxial leaf epidermis is the trichomes. Eight types of trichomes have been observed under SEM: simple-uniseriate, solitary, papillae, stellate, fused-stellate, stipitate-fasciculate, multiradiate and jellyfish-like. The first type, i.e. the simple-uniseriate trichome, is glandular, other six types, i.e. the solitary, papillae, stellate, fused-stellate, stipitate-fasciculate and multiradiate trichomes, are non-glandular, and the last one, i.e. the jellyfish-like trichome, may represent an intermediate type in structure between the glandular and the non-glandular trichomes. Papillae are very common in subgen. Cyclobalanopsis although in the Fagaceae they were previously reported only in a few species of Lithocarpus. The jellyfish-like trichome, observed only in Q. sichouensis, is reported in the Fagaceae for the first time. The evolutionary trends of trichomes are discussed. The trichomes seem to evolve toward the direction of increasing complexity in structure. Papillae are considered as the pedomorphism of trichomes. From papillae the solitary trichomes and then the stellate ones have evolved. The complicated trichome types with more arms may be produced from relatively simple ones with fewer arms by means of varions fusion: several stellate trichomes are fused to form the fused-stellate ones by basal fusion, and by the same way the fused-stellate ones to form the stipitate-fasciculate ones; the multiradiate trichomes are formed by the stellate ones, in which the arms emerge in a variety of seemingly random directions from a typically rounded common base. Based on the above results, it is considered that the trichome features are of important significance for a better understanding of the infrageneric division of Quercus and the phylogenetic relationships between this genus and the other genera in the Fagaceae. The very common presence of papillae in Quercus subgen. Cyclobalanopsis and in a few species of Lithocarpus may indicate that subgen. Cyclobalanopsis is a quite natural group on the one hand, and may have close relationship with Lithocarpus on the other hand. The thin-walled, jellyfish-like trichomes found in Q. sichouensis are in structure similar to the “glandular” peltate trichomes in Trigonobalanus, a genus generally regarded as the most primitive in subfam. Quercoideae, indicating that the affinity between subgen. Cyclobalanopsis and Trigonobalanus may becloser than that between subgen. Quercus and Trigonobalanus.     

栎属青冈亚属（壳斗科）的叶表皮研究

利用光镜和扫描电镜观察了栎属青冈亚属Quercus subgen. Cyclobalanopsis 48种植物的叶表皮,尤以对叶下表皮的毛被特征观察较为仔细。共观察到8种不同类型的叶表皮毛：单列毛、单毛、乳突、星状毛、溶和星状毛、具柄束毛、多出毛和水母状毛。其中乳突在青冈亚属中较常见,而在壳斗科其他属中仅在石栎属 Lithocarpus 少数种类中有报道;水母状毛首次在壳斗科中发现。毛被可能遵循以下的演化规律：乳突→单毛→星状毛;星状毛依照从简单→复杂的演化途径,分化出各种形态各异和结构复杂的毛系。初步讨论了毛被以及叶表皮其他特征(如毛基细胞和表皮细胞的形态、气孔的类型和密度等)的分类和系统学意义。     

Heterogeneity of stomatal density in the second wheat leaf

The stomatal density on abaxial epidermis of the second wheat(Triticum aestivum L.) leaf increased step by step from the leaf base to its tip. On adaxial epidermis, this increasing tendency was generally maintained, but close to both the base and the apex two maxima were determined. In the direction perpendicular to leaf length, maximum stomatal density was close to the edges, and minimum one was in the central part of the leaf blade.     

Differential regulation of trichome formation on the adaxial and abaxial leaf surfaces by gibberellins and photoperiod in Arabidopsis thaliana (L.) Heynh.

In wild-type (WT) Columbia and Landsberg erecta ecotypes of Arabidopsis thaliana (L.) Heynh., trichomes are present on the adaxial surfaces of all rosette leaves but are absent from the abaxial surfaces of the first-formed leaves. We have determined that both long-day (LD) photoperiod and gibberellin (GA) stimulate trichome formation. WT plants grown in LD conditions produce the first abaxial trichome on earlier leaves than plants grown in short-day (SD) conditions. Photoperiod sensitivity of abaxial trichome formation on WT plants develops gradually over time, reaching the maximum sensitivity about 24 d after germination. Application of gibberellic acid to WT plants growing in SD conditions accelerates the onset of abaxial trichomes. Conversely, application of 20 to 80 mg L-1 paclobutrazol, a GA biosynthesis inhibitor, to wild-type plants suppresses trichome initiation on the abaxial epidermis. The GA-deficient mutants ga1-5 and ga4-1 and the GA-insensitive mutant gai-1 exhibit delayed onset of abaxial trichomes when grown in LD conditions. The null mutant ga1-3 produces completely glabrous leaves when grown in SD conditions. Application of gibberellic acid to glabrous ga1-3 plants consistently induces earlier formation of trichomes on the adaxial epidermis than on the abaxial epidermis, demonstrating a difference between the adaxial and abaxial surfaces in their response to GA with regard to trichome formation.     

Morphological study on early ontogeny of theGinkgo leaf

The early ontogeny of theGinkgo leaf was studied by scanning electron microscope. The lamina of the leaf is derived from the protuberance of the adaxial side of a leaf buttress. The protuberance shows successive bifurcations. The second bifurcation occurs approximately at a right angle to the first. Thus, the abaxial surface of the leaf is derived from the outer surface of the protuberance and the adaxial is derived from the depressed inner surface of the protuberance formed by the first and second bifurcations. It is supposed that thick veins along both edges of the lamina are differentiated by an uneven dichotomous branching system of procambia caused by positional relationships: both edges of the lamina are derived from the parts of the protuberance just above a pair of procambia which come from the stem. On the shoot apex, there are a central cell group, which consists of only a few cells, and radially-arranged cell files. The leaf primordium is initiated from a sector area which consists of several files among radially-arranged cell files.     

Comparative Epidermal Ultrastructure of Cotton (Gossypium hirsutum L.) Leaf, Bract and Capsule Wall

Cotton leaves are more physiologically active than the bractand the capsule wall of the fruiting structures. To elucidatethe disparities in their physiological behaviour, epidermalcell density, stomatal index, stomatal size, trichome densityand type, and epicuticular wax ultrastructure of cotton leaf,bract and capsule wall were delineated using scanning electronmicroscopy (SEM). The epidermal cells of the outer periclinalwalls on both surfaces of the leaf and bract were raised andconvex. Conversely, the capsule wall epidermal cells were polygonalwith flat outer periclinal walls. The stomatal complex in theleaf and bract was paracytic, whereas in the capsule wall thestomatal complex was anomocytic. The adaxial and abaxial stomataof the leaf were coplanar to the epidermal surface, as opposedto the raised adaxial stomata on the bract. On the contrary,the stomata on the capsule wall surface appeared to be slightlysunken. Furthermore, the capsule wall stomata were larger thanthe stomata on either surface of both the leaf and the bract.The stomatal index was greater on the abaxial surfaces of theleaf and the bract (18.4 and 9.4, respectively) than their correspondingadaxial surfaces (14.4 and 4.7, respectively). Leaves had thehighest stomatal index followed by the bract and the capsulewall. The indumentum consisted of glandular and nonglandulartrichomes, the density of which was greater on the abaxial surfacesthan on the adaxial surfaces of the leaf and bract. The capsulewall indumentum lacked nonglandular trichomes. Epicuticularwax occurred in the form of striations. However, the striationpattern varied among the organs. This study clearly illustratesmorphological disparities in the epidermal features of leaf,bract and capsule wall, helping to explain their physiologicaldivergence. Copyright 2000 Annals of Botany Company Gossypium hirsutum, epicuticular wax, raised stomata, scanning electron microscopy, stomatal index, trichomes     

西安市常见绿化植物叶片润湿性能及其影响因素

利用接触角测定仪测定了西安市21种常见绿化植物叶片表面的接触角,探讨了叶片表面特性如蜡质、绒毛、气孔对接触角的影响。结果表明,植物叶片正背面、物种间的接触角差异均显著,叶片正面和背面接触角大小在40°～140°。接触角大小与变异系数呈负相关,可能由于接触角小的润湿叶片在不同的生境和位置下,受到环境条件的影响较大而出现大的变异;接触角较大的非润湿性叶片,环境物质持留时间较短,对叶片形态和组成影响较小,因而出现小的变异。植物叶片表面的接触角随蜡质含量的升高而增大。表皮蜡质去除后大部分叶片接触角明显降低,尤其是疏水性较强的银杏(Ginkgo biloba)、月季(Ro-sa chinensis)和紫叶小檗(Berberis thunbergii)。女贞(Ligustrum lucidum)正背面、加杨(Popu-lus canadensis)背面等亲水型的叶片蜡质去除后接触角反而增大。叶片绒毛的多少及其形态、分布方式对接触角具有重要的影响,不同的作用方式表现出润湿和不润湿的特征,人为将其去除可以增加叶片的润湿性。背面气孔密度与气孔长度、保卫细胞长度呈负相关;接触角则与气孔密度呈负相关,与气孔长度呈正相关。     

An investigation of the foliar trichomes of Tetradenia riparia (Hochst.) Codd [Lamiaceae]: An important medicinal plant of Southern Africa

The morphology and distribution of leaf trichomes of Tetradenia riparia were studied using light and scanning microscopy. Three morphologically distinct types of trichomes were observed on T. riparia leaf surfaces: glandular capitate (short and long stalked), peltate and non-glandular. The glandular and non-glandular trichomes were present in abundance on both the adaxial and abaxial surfaces. Young leaves were densely covered with trichomes; however, the density of trichomes decreases progressively with leaf maturity. This suggests that the trichomes are established early in leaf differentiation and their density decreases with leaf development and age.     

Comparative structure of the labellum in Ophrys fusca and O. lutea (Orchidaceae)

The morphology and anatomy of the labellar epidermal cells and the way in which they are arranged are described in an attempt to locate and characterize the osmophore in Ophrys fusca and O. lutea. The micromorphology of the labellum of these two species is similar. Four types of epidermal cells are present on the adaxial surface of the labellum. Long unicellular trichomes with straight tips cover the basal region of the labellum, whereas short unicellular trichomes with polygonal flattened bases form the reflective median speculum. The apical region of the labellum possesses a villous indumentum of long acuminate trichomes with bent or sinuate tips. Large smooth-walled, dome-shaped papillae occur on the margins and on the distal region of the abaxial surface of the labellum. These remarkable papillae have high polarity; the protoplasm at the apex of each cell contains several small vacuoles, while a prominent nucleus surrounded by numerous hypertrophied amyloplasts occurs at the opposite end of the cell. Positive reactions to Vogel's staining test and to Sudan black B enabled us to conclude that the osmophores of both species are composed of these peculiar secretory epidermal cells and by two or three subsecretory layers of parenchyma cells.     

DEVELOPMENT,STRUCTURE AND FUNCTION OF SECRETORY TRICHOMES IN PSYCHOTRIA BACTERIOPHILA (RUBIACEAE)

Mucilage-secreting dendroid trichomes develop from the adaxial epidermis of young stipules surrounding the shoot apex. Each trichome consists of a multicellular stalk from which radiate many branch cells. The trichome has no cuticle and the branch cell walls distally are loose cellulosic frameworks. Dictyosomes produce vesicles whose products are secreted through the plasma-lemma and cell wall. Enlarged portions of the ER are frequently associated with dictyosomes and may be part of the system for synthesis and transport of secretion products. Bacteria, which later occur in leaf nodules, are present in the mucilage surrounding trichomes and young leaves. The latter develop stomata through which the bacteria enter. As stipules and leaves grow out of the apical region, the secretory trichomes degenerate and are replaced by non-secretory ones.