Epidermal structure in the ligules of four species of the genus Poa L. (Poaceae)

The structure of the abaxial epidermis of the ligules of mid- and upper-culm leaves of the grasses Poa annua, P. nemoralis, P. pratensis and P. trivialis as seen by light microscopy is described. Amongst the structural elements found were long cells, short cells and unicellular prickle hairs over the surface, with unicellular hairs and papillate cells on the free edges of the ligules. Ligules of upper-culm leaves of some plants of P. trivialis L. contained vascular tissue with associated green (chlorenchyma) regions and stomata-like structures. The value of ligule characters in grass identification and classification is briefly discussed.     

ASSOCIATION OF MUCILAGE WITH THE LIGULE OF SEVERAL SPECIES OF SELAGINELLA

Mucilage bodies of protein and some polysaccharide were closely associated with ligules of Selaginella kraussiana (Kunze) A. Braun grown in the greenhouse, and Selaginella wallacei Hieron. and Selaginella oregana D.C. Eat. from natural habitats. With S. wallacei and S. oregana, several mucilage bodies were associated with some ligules. No mucilaginous material was associated with the ligules of Selaginella douglasii Hook, et Grev., Selaginella densa Rydb. or Selaginella apoda (L.) Fernald from natural habitats. When stem tips of S. kraussiana and S. douglasii were placed in sterile culture on a medium with sucrose, massive amounts of mucilage appeared around the ligules in the tight interstices of the stem tip leaves and extended out on the surface of the leaves. The mucilage from a cultured stem tip of S. kraussiana contained 0.034 μgm of protein, 14.6 μgm of apparent glucose equivalents as polysaccharide and no significant amounts of phenolics. In stem tips with tightly appressed leaves, either ligules or young leaves could have produced the mucilage; however, in more mature regions of the stem where internodal elongation had separated the leaves, mucilage coated only the apical portions of the ligules and was not associated with leaves. No other possible source of mucilage (bacteria or fungi) was detected among the leaves. When stem tips of S. wallacei, S. densa and S. apoda were placed in culture, mucilage was not produced. Since mucilage is closely associated with the ligule of some species of Selaginella growing in nature, the greenhouse, or tissue culture, the ligule may be glandular.     

FLORAL FLAVONOIDS AND ULTRAVIOLET PATTERNS IN VIGUIERA (COMPOSITAE)

Variation occurs among species of Viguiera series Viguiera for ultraviolet (uv) absorption/reflection patterns of ligules. Floral flavonoids that cause uv absorption occur in epidermal papillae. Flavonoids are further localized to the proximal portion of the ligule in the seven taxa that have only proximal uv absorption. Floral flavonoids involved in uv absorption consist of flavone, flavonol, and anthochlor (chalcone/aurone) glycosides. Quercetin 3-methyl ether glycosides characterize the ligules of 10 taxa occurring in Baja California, Mexico, and nearby areas, and these taxa appear to form one taxonomic group. The anthochlor pair, marein/maritimein, characterizes V. dentata, and the lack of ligule flavonoids distinguishes V. potosina from the remaining taxa. The presence of the anthochlor pair, marein/maritimein, only in V. dentata and the lack of ligule flavonoids in V. potosina concur with other data to indicate that these species are not correctly placed with each other or with the other species currently included in series Viguiera.     

LECLERCQIA COMPLEXA: EARLIEST LIGULATE LYCOPOD (MIDDLE DEVONIAN)

Our discovery of a ligule on Leclercqia complexa Banks, Bonamo and Grierson 1972 is the earliest occurrence of a ligulate lycopod in the fossil record. The ligule 1) occurs on a homosporous lycopod, differing with current concepts that the ligule is linked with the heterosporous condition; 2) is located on the leaf far distant from the attachment of leaf to stem, thus differing in position from any known ligulate lycopod, extinct or extant; 3) is comparable in morphology to ligules of extant lycopods, therefore providing no clues as to any earlier specialized function. These findings extend the enigma of the function of the ligule back in time, but necessitate a re-evaluation of the spatial relationship of the leaf and the ligule and of the link between heterospory and the ligule.     

VARIATION IN CALLOSE DEPOSITION IN THE LIGULES OF SEVEN SPECIES OF SELAGINELLA

The deposition of ligular callose, as defined and measured by fluorescence microscopy, was found to vary among different species of Selaginella. Species from dry, sunny habitats possessed greater quantities of callose in leaf ligules than did those species native to wet, shaded habitats.     

中国产3种水韭叶舌形态结构的比较研究

以中国产3种水韭的成熟叶片基部为实验材料,采用石蜡切片法,详细观察3种水韭的叶舌结构,建立各自叶舌的立体模型,探讨3种水韭间的亲缘关系,并比较了中国特产水韭与中亚及北美水韭的演化关系。结果显示:(1)3种水韭叶舌结构主要分为舌片、舌足和横带三部分。(2)中华水韭与东方水韭舌片均为三叉型,有主侧舌片之分,足角均为卵圆形,舌足与叶轴平行,而云贵水韭长鼻形,无主侧舌片之分,足角为三角形,舌足与叶轴有一定的角度。(3)中华水韭与云贵水韭的舌足侧面观为长椭圆形,东方水韭舌足侧面观为肾形。研究认为,中华水韭与东方水韭的亲缘关系较近,二者与云贵水韭亲缘关系较远;叶舌是水韭类的溯祖形态,其结构复杂,特征稳定,种间差异明显,应该作为水韭分类的重要依据。     

FLORAL NECTARIES AND TRICHOMES IN RELATION TO POLLINATION IN SOME SPECIES OF THEOBROMA AND HERRANIA (STERCULIACEAE)

Floral surfaces of two Theobroma spp. and one species of the allied genus Herrania (Sterculiaceae) were examined using scanning electron microscopy for reward system information of the pollinator-adaptation syndrome. These flowers are structurally complex and have sheltered reproductive parts indicating an elaborate insect-mediated pollination mechanism. Analysis of anther dehiscence, anthesis and pollen tube growth suggests a crepuscular pollination strategy. Suspected pollinators have been observed on floral parts of theobromas possessing stomate type nectaries and various shaped trichomes (blade of petal ligule and adaxial surface of petal hood). H. purpurea has elongate ligules devoid of stomates. Glandular clublike structures are found on the ovaries of all three spp. In numerous spp. of theobromas, except T. cacao, and two species of Herrania, a conspicuous ring of multicellular trichomes occurs at the adaxial base of the sepals. A possible function for the ring may be fragrance production because of its greatly reduced condition in the faintly scented T. cacao and the prevalence of pronounced fragrances in these other species.     

The prediction of leaf canopy expansion in the leek from a simple model dependent on primordial development

In an analysis of leaf development of leek plants grown in the field in 1988, successive leaves initiated, appeared (tip and ligule) and senesced at equal intervals of accumulated temperature/thermal time. These intervals corresponded to a plastochron of 92°C days and phyllochrons of 135 (tip) and 233 (ligule) °C days. The rate of appearance of ligules was exactly equal to the rate of leaf senescence, with the result that the number of fully-expanded leaves per plant remained constant at 1.4. These data, which were compatible with results from previous seasons, were used to develop a model of the interrelationships between primordium initiation at the shoot apex and subsequent events in the development of individual leaves. Primordium initiation is considered to be the primary controlling event in the life of a leaf, and the processes of tip appearance, ligule appearance and death can be predicted from knowledge of the number of primordia which have been initiated, without reference to the environment. A model of canopy expansion, based on the central role of the shoot apex, was developed using the temperature relations of primordium initiation and additional data on leaf expansion and leaf dimensions. Leaf area indices computed in this way provided a satisfactory simulation of the thermal-time course of leaf area index observed in a previous season, 1985.     

Structure and Function in the Grass Ligule: the Newly-initiated Ligule of Lolium temulentum L., a Light and Electron Microscope Study

Initiation and early development of the membranous ligule ofLolium temulentum L. was studied by light and transmission electronmicroscopy. The ligule appeared to be derived solely from theleaf adaxial epidermis and at this stage had a structure andultrastructure typical of a meristematic tissue. Cells of thefuture collar region of the leaf appear to be initiated at thesame time as the ligule. The siting of the ligule upon the leafis briefly discussed. Darnel, ligule (initiation), Lolium temulentum L., Poaceae, ultrastructure     

On the homology of the scales in Gunnera (Gunneraceae)

In the genus Gunnera , the species of subgenus Panke are unique in having large, triangular scales between the leaves on the rhizomes. The morphological significance of these scales has been extensively debated in the past. They have been interpreted as stipules, ligules, or cataphylls, with attempts made to identify homologous structures in representatives of the other subgenera. In the stoloniferous subgenus Misandra, the shoot apex is covered by a hoodlike structure, usually termed ochrea, generally considered homologous to the scales in Panke . Efforts to find similar structures in the remaining subgenera were unsuccessful. Due to the lack of a phylogenetic hypothesis, the homology of the scales has remained unresolved. In this study, the shoot apices of species from all subgenera were examined by light and scanning electron microscopy and the results interpreted in a phylogenetic context. The tip of the stolons in subgenera Pseudogunnera and Milligania carries two opposite cataphylls at the base of the new shoot. These are homologous to the ochrea in Misandra which is in turn homologous to the scales in Panke ; the latter are thus neither stipules nor ligules but cataphylls. The stems of subgenera Ostenigunnera and Gunnera have no similar structures. © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 142 , 301−308.     

Structure and Function in the Grass Ligule: Optical and Electron Microscopy of the Membranous Ligule of Lolium temulentum L.

Aspects of the structure and ultrastructure of the membranousligule of mature leaves of Lolium temulentum L. are described.In transverse section the ligule was lens-shaped and wedge-shapedin longitudinal section, 6 or 7 cells wide near the base and1 or 2 cells wide at the edges. Two uniseriate epidermes encloseda chlorenchymatous mesophyll tissue of varying thicknesses;both epidermes were continuous with the leaf adaxial epidermis.The cells comprising these three issues all appeared like typicalgrass epidermal long cells; elongate papillate cells were presentat the edges. No stomata, trichomes, intercellular spaces orvascular tissue were found in the ligule. A marked polarizationof ultrastructural complexity existed from the large-vacuolateabaxial epidermis to the ‘densely cytoplasmic’ small-vacuolateadaxial epidermis. Cells of the latter tissue contained numerousmitochondria, hypersecretory dictyosomes and abundant strandsof rough endoplasmic reticulum. Fluorescence microscopy providedevidence for the accumulation of a polysaccharide-containingmaterial within the periplasmic space next to the outer tangentialwall of adaxial epidermal cells. The ligule is considered tobe a highly organized and differentiated leaf organ with a pholosyntheticmesophyll and an adaxial epidermis active in the synthesis ofprotein and polysaccharide. Darnel, fluorescence microscopy, ligule, Lolium temulentum L., Poaceae, ultrastructure     

The Ligule of Isoetes lacustris: Ultrastructure, Mucilage Composition, and a Possible Pathway of Secretion

Mature ligules of Isoetes lacustris can be divided anatomicallyinto three ultrastructurally different regions. First, the basalregion contains large numbers of two types of protein bodies.Second, the cells of the sub-marginal region are characterizedby a well developed Golgi apparatus closely associated withthe tubular compartments of the endoplasmic reticulum. Third,the peripheral region consists of one to three layers of cellsshowing an extremely well developed rough asternal and smoothtubular endoplasmic reticulum (ER). The tubular ER compartmentsare frequently observed in close attachment to the plasmalemma.The outermost peripheral cells are covered with a mucilaginouslayer. The dry matter in the mucilage consists of 49 per centpolysaccharides and 22 per cent proteins. The polysaccharidefraction, analysed by ion exchange chromato-graphy, consistsmainly of glucose, arabinose, galactose and uronic acids. Theprotein fraction was analysed by SDS gel dectrophoresis andby high performance liquid-chromatographic separation of theamino acids. The analysis shows a protein pattern very similarto that of the peripheral ligule tissue. It is suggested, therefore,that the material of the external mucilage is secreted by theperipheral ligule cells. The secretional mechanism appears tobe a direct release of polysaccharides and proteins by the tubularcomponents of the ER. There is no indication of secretion viathe Golgi apparatus. Because of its high activity in proteinsynthesis and secretion, it is suggested that the ligule isa vestigial structure, which, in extinct genera, might havefunctioned as a digestive organ. Isoetes lacustris, endoplasmic reticulum, ligule, ultrastructure, Polysaccharide secretion, protein secretion     

Epidermal Structure in the Ligule of Rice (Oryza sativa L.)

The structure of the abaxial epidermis of the ligule of rice(Oryza sativa L.) as seen in the light and scanning electronmicroscope is described. Long cells, silica cells, cork cells,prickle hairs, microhairs and stomata-like structures were found.Reasons for considering these latter structures to be true stomataare given. Results of an X-ray analysis for silicon are presentedand the distribution of silica discussed in relation to itsfunctional role in the ligule. Oryza sativa L, rice, Gramineae, ligule, epidermis, scanning electron microscopy, X-ray analysis, silica, stomata     

Taxonomy of Selaginella: a study of characters, techniques, and classification in the Hong Kong species

This study of 11 Hung Kong species of Selaginella indicates that sufficient characters exist by which they may be conveniently, consistently, and conclusively distinguished, and implies that the same is true for the rest of the genus, which suffers from chronic taxonomic confusion. Characters assessed are: leaf size, shape, epidermal cell patterns and mesophyll structure; ligule shape; patterns seen in stem transections; sporophyll shape; microsporangium shape; sporangial distribution; and spore surface ornamentation. Of these, the shape of median leaves, the maximum size of lateral leaves and epidermal cell patterns are the most significant and distinctive foliar features. These, together with spore surface ornamentation, are the most useful for distinguishing species. Correlations between the shapes of ligules, sporophylls, microsporangia, and vascular bundles indicate natural subgeneric groups.
These studies suggest that there are two ways by which taxonomic investigations of Selaginella can be improved. First leaves and spores should be prepared in permanent mountant (e.g. Hoyer's solution) for accurate observation of size, shape, and epidermal patterns. Secondly, data should be collected on a wide range of features.     

Structure and Function of the Root Cap of Lolium temulentum L. (Poaceae): Parallels with the Ligule

Structure of the root cap ofLolium temulentum L. and of thechemistry of its secretory product were investigated with conventionaltransmission electron microscopy, zinc iodide–osmium tetroxideimpregnation, and use of protein and polysaccharide localizationtechniques at the ultrastructural level. Structural and functionalinformation concerning the root cap is compared with that previouslyobtained for the membranous ligule of this species. Similaritiesbetween root cap cells and adaxial epidermal cells of the liguleare described and discussed. It is concluded that both systemsare secretory and represent a similar response to the commonproblem of easing the passage of growing organs against thesoil or other plant parts. Darnel; Gramineae; grasses; ligule; Lolium temulentum L.; Poaceae; root cap; secretion; ultracytochemistry; ultrastructure; zinc iodide-osmium tetroxide impregnation     

Structure and function in the grass ligule: The membranous ligule ofLolium temulentum L. as a secretory organ

Summary Using phosphotungstic acid and periodic acid/thiocarbohydrazide/silver proteinate (Thiéry test) aspects of polysaccharide ultracytochemistry were studied in the membranous ligule ofLolium temulentum L. Staining results are presented for all three tissues-abaxial and adaxial epidermes and mesophyll-but discussed only for the epidermes. PTA- and PATAg-staining of the adaxial epidermis suggested synthesis of a conjugated polysaccharide material in this tissue, its accumulation in the periplasmic space and its subsequent secretion to the outside of the ligule via gaps in the cuticle. The ligule of this grass is considered to be a secretory organ.     

A New Species of Gigantochloa Kurz ex Munro (Poaceae: Bambusoideae) from Yunnan Province,China

A new species of Gigantochloa Kurz ex Munro, Gigantochloa callosa NH. Xia, Y. Zeng & RS. Lin was described and illustrated. It is differed from Gparvifolia (Brandis ex Gamble) TQ. Nguyen by its smaller size; culm sheath proper covered with both dark brown and silvery appressed hairs abaxially; ligules 2-3mm tall, entire; leaf ligules 2-3mm tall, entire; one side of the apex of leaf sheath developed into a 2-3mm long, thin, scale like callus.     

毛冠菊属舌片的微形态特征及其系统学意义

在扫描电镜下观察了毛冠菊属Nannoglottis 8种植物舌片的微形态特征,以探讨其颇有争议的系统位置。所有种类舌片近轴面的细胞为板状,长圆形;径向、切向壁直;外壁特征性突起,外壁中央为纵向皱纹,相邻细胞之间的纹饰无系统连接。这些特征表明毛冠菊属和紫菀族Astereae关系密切,但它在紫菀族内的系统位置可能比较特殊。这些结果和依据ITS序列推测的紫菀族系统发育的结果相吻合。文中还讨论了一些微形态性状的演化趋势以及微形态性状在毛冠菊属部分种类分类中的应用。