Epidermal Features of Leaves and Their Taxonomic Significance in Subfamily Ophiopogonoideae (Liliaceae)

This paper reports epidermal features of leaves in Ophiopogonoideae. Thirty-nine species and one variety (29 species, 1 variety in Ophiopogon, 6 species in Liriope, 4 species in Peliosanthes)were examined under scanning electron microscope. In addition, transections of stomatal apparatuses of six species (Ophiopogon: 3 species; Liriope: 2 species; Peliosanthes: 1 species) were made and examined under light microscope. The stomatal apparatus in Liriope, Ophiopogon and Peliosanthes is of the anomocytic type. These types of epidermal features of leaves in these genera are recognized: Cuticular processes type, No cuticular processes type and No stomatal band type. The cuticular processes type can be further divided into three patterns: Fibrillose, Massive and Wrinkled-massive. The taxonomic value of the epidermal features of leaves in Ophiopogonoideae is rather evident. (1)These epidermal features can be used to distinguish among those species of Ophiopogon, Liriope and Peliosanthes, even in their vegetative state; (2) The different patterns of cuticular processes are helpful to reasonable classification of some species in Ophiopogon, (3)They are of great value for recognizing some sections, (4) These epidermal features of leaves also provide evidence for further discussion on relationships among Ophiopogon, Liriope, and Peliosanthes. The evolutionary trend of the epidermal features of leaves in Ophiopogonoideae is No stomatal band type→No cuticular process type(stomatal band)→Cuticular process type (stomatal band). According to the epidermal features of leaves, flowers and fruits, Ophiopogon, Liriope and Peliosanthes are closely related, forming a subfamily Ophiopogonoideae. Ophiopogon is more close to Liriope than to Peliosanthes, and they should be grouped into the same tribe-Ophiopogoneae. Liriope seems to be more primitive than Ophiopogon. Peliosanthes, which constitutes another tribe of its ownPeliosantheae, is more advanced than Ophiopogon and Liriope, and it might have beenderived from its ancestor early.     

Pollen Morphology and Generic Phylogenetic Relationships in Ophiopogonoideae (Liliaceae)

In the present work, pollen grains of 3 species of Liriope, 24 species of Ophiopogon and 2 species of Peliosanthes were examined under scanning electron microscope, Among them 4 species were also observed under transmission electron microscope. The observation (Table 3)shows that Liriope and Ophiopogon distinctly differ from Peliosanthes in the exine ornamentation and structure, They may be divided into two types: 1. Rugulate-perforate, ektexine with perforate tectum in Liriope and Ophiopogon. 2. Verrucate, Verracae unequal in size, ektexine intectate in Peliosanthes. Pollen morphology shows the close affinity between Liriope and Ophiopogon, but they are very far from Peliosanthes. The correlation between pollen and gross morphology in Liriope, Ophiopogon and Peliosanthes are stated and their evolutionaly trends are discussed in this paper. The pollen characters support the placement of Liriope and Ophiopogon in one tribe—Ophiopogoneae, and Peliosanthes in another tribe—Peliosantheae. Peliosanthes is more advancedthan Liriope and Ophiopogon.     

Systematics of Ruscaceae/Convallariaceae: a combined morphological and molecular investigation

This paper discusses the intrafamilial systematics of Ruscaccae sensu lata by means of a combined molecular-morphological analysis. Ruscaceae sensu lata (including former Con-vallariaceae. Dracaenaceae and Nolinaccac, as well as Comospermum and Eriospermum ) represent a well-supported clade in other molecular analyses and are further linked by the absence of phytomclan in the seed coat. Within this clade there is an unusual lack of sequence divergence for such a morphologically varied group of taxa, but the combined morphological-molecular analysis indicates some groupings: (1) Eriospermum , (2) Comospermum , (3) nolinoids (former Nolinaceac) plus Ophiopogon and Liriope (Ophiopogonae, excluding Peliosanthes ), (4) dracaenoids (former Dracaenaceae), (5; Polygonatae and (6) a clade comprising Convallarieac (including Aspidistra ) and the ruscoids (Ruscaccae sensu stricto ) plus Peliosanthes. In the morphological analysis Peliosanthes is embedded in Convallarieae, and in the molecular analysis it is sister to all other Ruscaceae except Eriospermum: in neither case does it fall with Ophiopogon and Liriope (Ophiopogonae), with which it was traditionally placed. Peliosanthes has several characters in common with some Convallarieae, e.g. filaments fused into a column surrounding the gynoecium, and a thick, fleshy corona with the stigma more or less filling the narrow floral opening. On the other hand, there arc also some notable differences, such as in karyotype, leaf anatomy, pollen morphology and nectaries. Since there is a considerable range of morphological variation in the taxon currently designated as Peliosanthes. more taxonomic work is needed to establish the monophyly of this genus before there can be confidence in its relationships within Ruscaccae sensu lata.     

The leaf internal morphology and ultrastructure of Zostera muelleri irmisch ex aschers. (Zosteraceae): a comparative study of the intertidal and subtidal forms

The leaf anatomy, histochemistry and ultrastructure of the intertidal and subtidal seagrass Zostera muelleri Irmish ex Aschers. from Westernport Bay, Victoria were studied. Unusual anatomical and ultrastructural features are compared with other seagrasses and their functional significance is assessed. Subcuticular cavities are present in the young blade, but not observed in the older blade nor young and old leaf sheath. Wall ingrowths occur in the blade epidermal cells particularly on the inner tangential walls and the lower portions of the radial walls. Plasmodesmata are present between adjacent epidermal cells and between the epidermal and mesophyll cells, suggesting that solutes could transfer between these tissues both symplastically and apoplastically. Each leaf has three longitudinally aligned vascular bundles, each of which comprises a single xylem element isolated from the phloem tissue. The phloem consists of nacreous-walled sieve elements accompanied by phloem parenchyma cells which also process wall ingrowths. The xylem walls are completely hydrolysed and the middle lamella borders directly on the xylem lumen. Leaves have prominent air lacunae bisected transversely by septa at regular intervals along their length. Each septum consists of a file of small parenchyma cells with wall protuberances projecting into intercellular space. There are no major structural differences between the subtidal and intertidal plants, but the former have larger leaves and more leaves per shoot than the latter. In addition, a network of unusual reticulated fungal hyphae is present in the leaf intercellular spaces of the subtidal form and this network may facilitate solute transfer in these plants.     

Anatomical characterization of Strobilanthes (Acanthaceae) species from the northern Western Ghats of India and its implication for identification at vegetative state

A comprehensive study of stem, leaf and petiole anatomy of 10 species of Strobilanthes from northern Western Ghats of India was carried out to identify characteristics which would enable species identification when flowering material is unavailable. In Strobilanthes, some species bloom annually, others are plietesials, i.e. they grow without blooming for several years and then produce huge quantities of flowers, release seeds and die. Therefore, alternative methods, such as anatomical characters, are essential to distinguish Strobilanthes species in their vegetative stage. We collected ten species of Strobilanthes for anatomical characterization. Under the bright‐field microscope, stem cross‐sections of different species were found to be undulate, quadrangular, quadrangular‐winged or terete. Study of the stem revealed a distinct outer and inner cortex, the distribution of cystoliths (CaCO₃ crystals), raphides (CaC₂H₂O₅ crystals) and sclereids which varied from species to species. Study of leaf anatomy showed structural variation and vascular bundle shapes that differed between the species. Leaf epidermal characters under light and scanning electron microscopy exhibited variation in characters such as stomatal index, stomatal length and width, stomatal type and presence of glandular and non‐glandular trichomes. The petiole anatomy was species‐specific, especially with respect to vascular bundle structure and the distribution of structures such as sclereids, cystoliths, sphaeraphides and tannin cells varied. Hence, unique anatomical features of the stem, leaf and petiole could be used as taxonomic characters to identify Strobilanthes species in a vegetative state.     

Leaf and twig anatomy of Eriope, a xeromorphic genus of Labiatae

The leaf and twig anatomy of 25 species of the genus Eriope were studied. The twig anatomy is very uniform apart from the level of formation of early layers of cork. Leaf anatomy shows considerable variation between the species, and this is correlated to some extent with the extreme habit range from trees to woody herbs. Characters of the lamina that show variation are: trichome type and frequency, cuticular markings, leaf dorsiventral or isobilateral, presence of adaxial stomata, presence of a hypodermis, number of layers of adaxial palisade mesophyll cells, occurrence of large bundles of phloem fibres at main veins, type of areolation and marginal venation. Petiole vasculature is simple and generally with either four distinct vascular bundles or two vascular arcs. The most xeromorphic species are usually woody herbs or sub-shrubs, and tend to have thick, isobilateral leaves with large bundles of phloem fibres and few hairs, or strongly dorsiventral leaves with a hypodermis and stomata in deep abaxial hair-lined depressions. The correlation of xeromorphic characters with environmental conditions is discussed. Leaf anatomy is of limited value in elucidating relationships within the genus.     

星星草营养器官适应盐胁迫的结构特征

采用0．6％Na2CO3胁迫处理星星草[Puccinellia tenuiflora（Turcz．）Scribn．et Merr．]幼苗,光镜和电镜观察其根和叶的显微和超微结构。结果表明,星星草根的表皮向外突出形成密集的根毛;外皮层由1～2层细胞组成,排列较紧密;中皮层薄壁细胞排列疏松,形成发达的通气组织;内皮层呈典型的五面加厚;中柱鞘排列紧密,其壁加厚;初生木质部与初生韧皮部相间排列,初生木质部为5～7原型,中央为后生木质部导管,无髓存在。叶的表皮有表皮毛和丰富的蜡质层;叶上表皮泡状细胞数目较少,且深陷;气孔下陷,其下有较大的气室;叶脉有大、中、小3种维管束,大、中型维管束为C3型,小型维管束为C4型。星星草可能是介于C3和C4植物之间的类型,具有耐盐碱及耐干旱特征。     

Histopathology of Oil Palm Seedlings Infected by, Pathogenic Fusarium oxysporum Isolates from Africa

Oil palm seedlings (one-leaf stage) grown from Malaysian seed were inoculated with six African isolates of Fusarium oxysporum f. sp. elaeidis and one African isolate of Fusarium oxysporum var. redolens. All the isolates induced similar symptoms and anatomical responses in the inoculated palms. The pathogen invaded the root, bulb, leaf bases and leaves in that sequence. In the root and bulb, infection resulted in plugging of xylem elements with mycelium, conidia, tyloses and gums, disintegration and plugging of phloem tissues, disintegration of plugged vascular elements forming gaps and cavities, and collapse of cortical tissues leading to the isolation of the infected vascular tissues. Formation of hypertrophic or hyperplastic cells in the xylem parenchyma was not observed. Anatomical changes in the infected leaf bases and petioles were similar to those occurring in the infected bulb; except that tyloses were formed less frequently. Generally there was no sign of the pathogen in the wilted leaf blade, but the epidermal, hypodermal and mesophyll cells appeared shrunken or to have collapsed. No hyphae or anatomical changes were observed m the tissues of symptomless inoculated palms.     

Comparative anatomy of the young stem,node, and leaf of Bonnetiaceae,including observations on a foliar endodermis

A comprehensive study of the nodal and leaf anatomy of Bonnetiaceae was completed in order to provide evidence for evaluation in relation to systematics. Nodal anatomy is trilacunar, three-trace or unilacunar, one-trace. Basic leaf anatomical features of the family include: complete or incomplete medullated vascular cylinder in petiole; paracytic mature stomata with encircling ridges; large mucilaginous cells in the adaxial surface of mesophyll; periclinal divisions in upper surface layers; and discrete patches of phloem within the vascular bundles. Especially noteworthy is the presence in some genera of foliar vascular bundles enveloped by a sheath composed of two concentric regions, i.e., an inner region consisting of multiple layers of fibers and an outer specialized endodermis composed of thin-walled cells with Casparian strips. Leaves are variable with respect to lamina and cuticle thickness, relative amount and number of palisade and spongy layers, venation of lamina, and the presence or absence of sclereids and crystals in the mesophyll. A major feature in the evolution of Bonnetiaceae is development of a highly divergent, essentially parallel, leaf venation that is superficially similar to that of some monocotyledons and apparently unique among dicotyledons. Foliar anatomy provides important characters for the recognition of subgroups within Bonnetiaceae and is consistent with the segregation of Bonnetiaceae from Theaceae.     

The systematic position ofAsplenium cardiophyllum (Aspleniaceae)

Asplenium cardiophyllum is a morphologically unusual species with simple leaves and anastomosing venation, and is often placed in the segregate genusBoniniella. To determine its systematic position, character comparisons were made of vascular anatomy, raphides in leaf epidermis, chromosome number and perispore of this species and those ofAsplenium sect.Hymenasplenium. Asplenium cardiophyllum conforms with sect.Hymenasplenium in its dorsiventral dictyostele, the presence of raphides, a chromosome number of 2n=156 (x=39), and lophate peristore with spinulate projections on the lumina. We therefore propose to includeA. cardiophyllum in that section. Dedicated to the memory of the late Professor Kunio Mitui.     

六种苏铁属植物的羽片比较解剖学研究

研究了锈毛苏铁（Cycas ferruginea),石山苏铁（C.miquelii),四川苏铁（C.szchuanensis),海南苏铁（C.hainanensis),仙湖苏铁（C.fairylakea)和贵州苏铁（C.guizhouensis)等六种苏铁属植物羽片的比较解剖学,结果显示锈毛苏铁与石山苏铁在下皮层厚壁细胞,海绵组织中含晶细胞,中脉隆起,叶缘形态,韧皮部形态及分泌道的有无等特征上具有较明显的差异,四川苏铁与海南苏铁和仙湖苏铁非常相近;海南苏铁与仙湖苏铁基本一致;贵州苏铁在中脉隆起,叶缘形态,韧皮部形态等方面与四川苏铁,海南苏铁及仙湖苏铁三者有差异。支持将锈毛苏铁,石山苏铁独立为种的观点,并认为海南苏铁与仙湖苏铁不能区分,四川苏铁与海南苏铁和仙湖苏铁具有较近的种系关系,贵州苏铁与其它种的种系关系相对较远。     

Comparative and functional leaf anatomy of selected Alstroemeriaceae of mainly Chilean origin

The resupinate leaves of 16 species of Alstroemeriaceae were examined using light- and scanning electron microscopy. The leaf anatomy is described for all of the species, that of the petiole and stem for selected species. The mesophyll consists of chlorenchyma and includes idioblasts that contain raphides. Dorsiventral, isolateral or isobilateral leaf types were observed. Petioles are always isolateral. Two epidermal types are commonly observed: type I contains jigsaw puzzle-like intercostal cells and longitudinally elongated cells above the veins; type II contains only longitudinally elongated cells, usually longer, above the veins. Some species have an epidermis which differs from the main types. All species show adaptation to resupination by having an inverted anatomy. Due to the twist of the leaves, adaxial indicates the lower surface and abaxial the upper. Stomata are found on both surfaces. Palisade cells, when they occur, are always on the abaxial surface. Most species are mesomorphic in spite of the dry environments in which they grow; a few show xeromorphic features. The leaves are compared and discussed with relation to ecological conditions.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 261−272.     

Heterogeneity of silica and glycan-epitope distribution in epidermal idioblast cell walls in Adiantum raddianum laminae

Laminae of Adiantum raddianum Presl., a fern belonging to the family Pteridaceae, are characterised by the presence of epidermal fibre-like cells under the vascular bundles. These cells were thought to contain silica bodies, but their thickened walls leave no space for intracellular silica suggesting it may actually be deposited within their walls. Using advanced electron microscopy in conjunction with energy dispersive X-ray microanalysis we showed the presence of silica in the cell walls of the fibre-like idioblasts. However, it was specifically localised to the outer layers of the periclinal wall facing the leaf surface, with the thick secondary wall being devoid of silica. Immunocytochemical experiments were performed to ascertain the respective localisation of silica deposition and glycan polymers. Epitopes characteristic for pectic homogalacturonan and the hemicelluloses xyloglucan and mannan were detected in most epidermal walls, including the silica-rich cell wall layers. The monoclonal antibody, LM6, raised against pectic arabinan, labelled the silica-rich primary wall of the epidermal fibre-like cells and the guard cell walls, which were also shown to contain silica. We hypothesise that the silicified outer wall layers of the epidermal fibre-like cells support the lamina during cell expansion prior to secondary wall formation. This implies that silicification does not impede cell elongation. Although our results suggest that pectic arabinan may be implicated in silica deposition, further detailed analyses are needed to confirm this. The combinatorial approach presented here, which allows correlative screening and in situ localisation of silicon and cell wall polysaccharide distribution, shows great potential for future studies.     

白鲜营养器官解剖结构及其与白鲜碱的积累关系

应用植物解剖学、组织化学及植物化学方法对白鲜营养器官根、茎、叶的结构及其生物碱的积累进行了研究。结果显示:(1)白鲜根的次生结构以及茎和叶的结构类似一般双子叶植物;白鲜多年生根主要由周皮、次生韧皮部、维管形成层以及次生木质部组成,根次生韧皮部中可见大量的淀粉、草酸钙簇晶、韧皮纤维以及油细胞;茎由表皮、皮层、维管组织和髓组成;叶由表皮、栅栏组织、海绵组织和叶脉组成;在茎和叶初生韧皮部的位置均分布有韧皮纤维,在叶表皮上分布有头状腺毛和非腺毛;在茎和叶紧贴表皮处分布有分泌囊。(2)组织化学分析结果显示:在白鲜多年生根中,生物碱类物质主要分布在周皮、次生韧皮部、维管形成层和木薄壁细胞中;在茎中,生物碱主要分布在表皮、皮层、韧皮部、木薄壁细胞及髓周围薄壁细胞中;在叶中,生物碱主要分布在表皮细胞、叶肉组织和维管组织的薄壁细胞;此外在分泌囊和头状腺毛中亦含有生物碱类物质。(3)植物化学结果显示,秦岭产白鲜根皮/白鲜皮、根木质部、茎和叶中白鲜碱含量分别为0.041%、0.012%、0.004%和0.002%,其中木质部中白鲜碱含量和其他部分地区白鲜皮中白鲜碱含量类似。研究表明,在秦岭产白鲜营养器官中,除根皮/白鲜皮外,在根木质部亦含有大量的白鲜碱,且在茎和叶中亦含有一定的白鲜碱,具有潜在的开发利用价值。     

六种苏铁属植物的羽片比较解剖学研究

研究了锈毛苏铁（Cycas ferruginea）、石山苏铁（C. miquelii）、四川苏铁（C. szechuanensis）、海南苏铁（C. hainanensis）、仙湖苏铁（C. fairylakea）和贵州苏铁（C. guizhouensis）等六种苏铁属植物羽片的比较解剖学,结果显示锈毛苏铁与石山苏铁在下皮层厚壁细胞、海绵组织中含晶细胞、中脉隆起、叶缘形态、韧皮部形态及分泌道的有无等特征上具有较明显的差异;四川苏铁与海南苏铁和仙湖苏铁非常相近;海南苏铁与仙湖苏铁基本一致;贵州苏铁在中脉隆起、叶缘形态、韧皮部形态等方面与四川苏铁、海南苏铁及仙湖苏铁三者有差异。支持将锈毛苏铁、石山苏铁独立为种的观点,并认为海 南苏铁与仙湖苏铁不能区分。四川苏铁与海南苏铁和仙湖苏铁具有较近的种系关系,贵州苏 铁与其它种的种系关系相对较远。     

Leaf anatomical studies of Eugenia L. (Myrtaceae) species from the Malay Peninsula

KHATIJAH, H. H., CUTLER, D. F. & MOORE, D. M., 1992. Leaf anatomical studies of Eugenia L. species from the Malay Peninsula. The leaf and petiole anatomy of 25 Malaysian species of Eugenia L. sensu lato was investigated to determine the value of vegetative anatomy for the identification of non-flowering material. Twelve characters proved useful for the identification of species: type of stomata; shape of leaves in T.S.; shape of midrib bundle; sclerenchyma sheath; cutinization of outer epidermal wall; presence or absence of sclereids, idioblasts, hypodermis, columnar epidermal cells, solitary crystals; number of palisade layers and the shape of vascular strand, sclerenchyma sheath and sclereids in the parenchyma of the petiole. Using these characters an anatomical key was constructed to assist identification of the species. Only the closely related species E. stapfiana and E. nilidula could not be separated on anatomical characters.
The anatomical data support the treatment of Henderson (1949) of section Acmena but not of sections Cleistocalyx and Fissicalyx , which fit better in section Syzygium.     

地枫皮营养器官解剖结构特征及其叶片结构的生态适应性

为了明确地枫皮的结构特征及其在石灰岩山顶和山腰疏林间两种环境下生长的叶片解剖结构的差异,本研究采用石蜡切片和半薄切片对地枫皮营养器官进行解剖观察并评价了叶片结构对不同生态环境的响应。结果表明:地枫皮根中次生维管组织发达,木射线和韧皮射线明显。老茎的次生构造中,皮层贮藏物质丰富,内有大的石细胞群,韧皮射线和木射线明显;而髓细胞内含有大量晶簇和少量单晶。在叶片横切面观上,叶为异面叶,表皮细胞一层,上表皮无气孔分布,主脉中薄壁细胞中分散有石细胞。叶片解剖结构显示,随海拔高度上升,地枫皮趋向于旱生植物的特点,其主要表现在叶片表皮细胞外壁角质层加厚,栅海比增加、海绵组织排列由紧密变疏松。另外,根皮、茎皮和叶肉中都分布有大量油细胞。     

Leaf anatomy and relationships of Dietes (Iridaceae)

The leaf anatomy of Dietes is described, including all 6 species. Characters support the morphological evidence that Dietes is a distinct genus and related to other Iridoideae. Certain features, notably the epidermal structure and sheath vasculature, distinguish it from other Iridaceous genera, although the sheath vasculature indicates a relationship with some New World Iridoideae. The leaf margin type is similar to that of other Old World Iridoideae. Many leaf anatomical characters in Dietes are associated with xeromorphy. Leaf anatomy supports the division of Dietes into 2 subgenera.