STIGMA,STYLE, AND OBTURATOR OF ORNITHOGALUM CAUDATUM (LILIACEAE) AND THEIR FUNCTION IN THE REPRODUCTIVE PROCESS

Transmitting tissue in Ornithogalum is divided into three regions corresponding to classical divisions of the gynoecium: stigma, style, and ovary. The stigma differentiates from epidermal cells of the stylar apex. These cells form the stigmal papillae and have dense cytoplasm with abundant ER and lipid bodies. Papillae have walls with small transfer-ingrowths. At floral receptivity, papillae secrete a small amount of surface exudate. Epidermal cells of the style contain numerous spherosomes and have thin filaments of cytoplasm traversing the central vacuole. The stylar cortex is composed of 3-6 layers of parenchyma cells which contain numerous spherosomes and often have secondary vacuoles. Vascular tissue in the style consists of one collateral bundle in each lobe. Cells of the epidermal layer lining the stylar canal are secretory. They are initially vacuolate but fill progressively with dense cytoplasm as their secretory activity increases. Secretory activity occurs in three phases, each characterized by a particular organelle population and secretory product. At anthesis, the canal is filled with an exudate consisting of carbohydrate, protein, and lipid. In the ovary, the obturator differentiates from cells at the base of the funiculus and the tip of the carpel margins. It forms a pad of tissue which covers most of the former placenta. The obturator is secretory and produces a surface exudate. We believe our observations on Ornithogalum support the hypothesis that all transmitting tissue is of the same morphological origin and that it provides nutritive and chemotropic factors for pollen tube growth.     

Flower palate structure of the aquatic bladderworts <Emphasis Type="Italic">Utricularia bremii</Emphasis> Heer and <Emphasis Type="Italic">U. minor</Emphasis> L. from section <Emphasis Type="Italic">Utricularia</Emphasis> (Lentibulariaceae)

There is an enormous diversity in the structure of the flower palate of the carnivorous rootless genus Utricularia. This study aims to examine the structure of the palates in Utricularia bremii Heer and U. minor L of the Utricularia sect. Utricularia, which have a glandular palate type. In both species, the palate has only one type of glandular trichomes. Because of the occurrence of cell wall ingrowths in its glandular cells, any exudation may be transported via eccrinous secretion. It was proposed that the palate trichomes of the examined species act as scent glands and that the palate may play a role as an unguentarium. Both U. bremii and U. minor are of an open flower type. Thus, U. bremii and U. minor flowers can be penetrated by small, weak insects, which then easily have access to their generative structure. Small Hymenoptera (member of families Mymaridae and Braconidae) were observed as flower visitors of the male-sterile species Utricularia bremii.     

Pollination biology of mass flowering terrestrial utricularia species (lentibulariaceae) in the Indian Western Ghats

The pollination biology of three mass flowering Utricularia species of the Indian Western Ghats, U. albocaerulea, U. purpurascens, and U. reticulata, was studied for the first time by extensive observation of flower visitors, pollination experiments, and nectar analyses. The ephemerality of the Utricularia habitats on lateritic plateaus, weather conditions adverse to insects, lack of observations of flower visitors to other Utricularia spp., and the predominance of at least facultative autogamy in the few Utricularia species studied so far suggested that an autogamous breeding system is the common case in the genus. In contrast, we showed that the studied populations are incapable of autonomous selfing, or that it is an event of negligible rarity, although P/O was similarily low as in autogamous species investigated by other authors. In all three species the spatial arrangement of the reproductive organs makes an insect vector necessary for pollen transfer between and within flowers. However, U. purpurascens and U. reticulata are highly self-compatible, which allows for visitor-mediated auto-selfing and geitonogamy on inflorescence and clone level. Floral nectar is present in extremely small volumes in all three species, but sugar concentrations are high. More than 50 species of bees, butterflies, moths, hawk moths, and dipterans were observed to visit the flowers, and flower morphology facilitated pollination by all observed visitors. The results are discussed in the context of the phenological characteristics of the studied species, especially the phenomenon of mass flowering, and the environmental conditions of their habitats.     

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。     

Recent progress in understanding the evolution of carnivorous lentibulariaceae (lamiales)

Carnivorous plants have emerged as model systems for addressing many ecological and evolutionary questions, and since Lentibulariaceae comprise more than half of all known carnivorous species (325 spp.), they are of particular interest. Studies using various molecular markers have established that Lentibulariaceae and their three genera are monophyletic with Pinguicula being sister to a Genlisea-Utricularia-clade, while the closest relatives of the family remain uncertain. Character states of the carnivorous syndrome in related proto-carnivorous lamialean families apparently emerged independently. In Utricularia, the terrestrial habit has been reconstructed as plesiomorphic, and an extension of subgenus Polypompholyx is warranted. In the protozoan-attracting Genlisea, subgenus Tayloria is revealed as basal lineage. In Pinguicula, the six major lineages found reflect radiations in clearly defined geographic regions, whereas most previously recognized subgeneric taxa are non-monophyletic. Genlisea and Utricularia exhibit substitutional rates that rank among the highest in angiosperms for the molecular markers analyzed. One possible explanation for this lies in selective constraints on a wide range of genomic regions that may have been lowered due to the use of an alternative mode of acquiring nutrients.     

Studies on Two Palaeozoic Seeds from China

This paper reports two species of the seed order Cardiocarpales: Cardiocarpus (?) minor (Wang) emend and Diplotesta sp. which were found in coal balls. The specimens were collected from Taiyan Formation of Upper Carboniferous near Xiedao Village, West Mountain, 'Taiyan City, Shanxi Province and Wangjiazhai Formation of Upper Permian of Wangjiazhai Mine of Shuicheng district, Guizhou Province, China. The two taxa are described and their taxonomy is discussed in detail. The diagnosis of the species (Cardiocarpus (?) minor (Wang) emend) is amended as follows: Seeds bilaterally symmetric and thinner cardioformed, appoximatly 4–6 mm long, 3–4 mm wide in the primary section and less than 2 mm thick in the secondary section. Integument differentiated into three zones. Outer sarcotesta delimkted into four layers: 1) carbonized cuticle partly preserved; 2) epidermis of thin-walled parenchyma 1–2 rows of small cells; 3) hypodermis of secretory tissue containing large cells, uniform in thickness; and 4) parenchyma layer consisting of thin-walled cells in size varying from 20 to 50 μm middle sclerotesta of vertically arranged fibers is bipartite: outer fibers being smaller in diameter and possessing thicker walls than inner fibers. Sclerotesta 80 μm in thickness commissural laterally to form two lateral ribs. Inner endotesta single rowed of parenchymatous cells. Nucellus free except at base which is attached to the integument by narrow nucellar platform. Pollen chamber not observed. The double vascular system includes integumentary strand beneath secretory layer of sarcotesta and nucellar strand which forms a tracheid mentle; the origin of integumentry and nucellar strands unknown. In addition a computerized three-dimensional reconstuction techniqne is utilized to reconstruct membrane like structure of Diplotesta sp. It is suggested that Cardiocarpus (?) minor is different from other species of the genus Cardiocarpus in the distribution of nucellar vascular tissues. This feature is thonght to be one of the important characters of bilaterally symmetrical seeds in a taxonomic view point. Therefore Cardiocarpus minor wang probally represents a menber of another new seed genus, althongh this species remains in the genus Cardiocarpus.     

Floral nectary and osmophore of <Emphasis Type="Italic">Epipactis helleborine</Emphasis> (L.) Crantz (Orchidaceae)

The analysis of flowers collected at different stages of anthesis provides strong evidence to conclude that the shell-shaped hypochile and the knobs of epichile form a nectary. The scent comes from the aromatic constituents of nectar and the epichile tissue and the apices of all tepals (osmophores). The comparison between pollinated and unpollinated flowers revealed that the anthesis of unpollinated flowers lasted up to the 16th day. The nectariferous secretory cells formed single-layered epidermis and several layers of underlying parenchyma built by small, isodiametric cells with thin walls and dense cytoplasm, relatively large nuclei, supplied by collateral vascular bundles. During the floral lifespan, the residues of secreted material were higher on the hypochile cells. The lipoid-carbohydrate material and lipid globules in the cell walls and in the cytoplasm were localised. The abundance of starch grains was observed at the beginning of anthesis and their gradual reduction during the flower lifespan. At the end of anthesis in unpollinated flowers, the lipoid-carbohydrate-phenolic materials have been demonstrated. The phenolic material was the same as in plastoglobuli. The features such as irregular plasmalemma, the secretory vesicles that fuse with it, fully developed dictyosomes, numerous profiles of ER indicate vesicle-mediated process of secretion. The substances could be transported by vesicles to the periplasmic space via granulocrine secretion and then to the external surface. Both micro-channels and slightly developed periplasmic space were visible in the hypochile epidermis. This is the first time for anatomical survey of secretory tissue in pollinated and unpollinated flowers of E. helleborine.     

Comparative Anatomy of the Secondary Phloem of the Stem in Cephalotaxus

The family Cephalotaxaceae contains only one genus, Cephalotaxus, with 9 species and some varieties, mostly endemic to China. The present paper deals with the comparative anatomy of secondary phloem of stems in 4 species and I cultivar, of this genus under light microscope and scanning electron microscope. The main results are as follows: (1) The structure of secondary phloem is rather uniform in the various species of this genus. In cross section, sieve cells, phloem parenchyma cells form continuous tangential rows of one cell in width respectively, which occur alternately. Sclerenchyma cells also form continuous tangential rows, each with a radial width of 1-4 cells. The interval between the rows is rather wide. (2) The type of phloem fiber and the quantity of sclereids can be served as the characters for identification of species and the evidence for the separation of two sections of Cephalotaxus, namely: Cephalotaxus and Pectinatae. (3) The secondary phloem of this genus contains more or less crystalliferous parenchyma cells, in the inner tangential walls of which calcium oxalate crystals are embedded. So far this character has not been reported in the other families and the genera of conifers. Our conclusion, therefore, agrees with the opinion that Cephalotaxaceae is a natural taxon and includes only one genus, Cephalotaxus.     

Female germ unit in Genlisea and Utricularia, with remarks about the evolution of the extra-ovular female gametophyte in members of Lentibulariaceae

Lentibulariaceae is the largest family among carnivorous plants which displays not only an unusual morphology and anatomy but also the special evolution of its embryological characteristics. It has previously been reported by authors that Utricularia species lack a filiform apparatus in the synergids. The main purposes of this study were to determine whether a filiform apparatus occurs in the synergids of Utricularia and its sister genus Genlisea, and to compare the female germ unit in these genera. The present studies clearly show that synergids in both genera possess a filiform apparatus; however, it seems that Utricularia quelchii synergids have a simpler structure compared to Genlisea aurea and other typical angiosperms. The synergids are located at the terminal position in the embryo sacs of Pinguicula, Genlisea and were probably also located in that position in common Utricularia ancestor. This ancestral characteristic still occurs in some species from the Bivalvaria subgenus. An embryo sac, which grows out beyond the limit of the integument and has contact with nutritive tissue, appeared independently in different Utricularia lineages and as a consequence of this, the egg apparatus changes position from apical to lateral.     

Functional utrastructure of Genlisea (Lentibulariaceae) digestive hairs

BACKGROUND AND AIMS: Digestive structures of carnivorous plants produce external digestive enzymes, and play the main role in absorption. In Lentibulariaceae, the ultrastructure of digestive hairs has been examined in some detail in Pinguicula and Utricularia, but the sessile digestive hairs of Genlisea have received very little attention so far. The aim of this study was to fill this gap by expanding their morphological, anatomical and histochemical characterization. METHODS: Several imaging techniques were used, including light, confocal and electron microscopy, to reveal the structure and function of the secretory hairs of Genlisea traps. This report demonstrates the application of cryo-SEM for fast imaging of whole, physically fixed plant secretory structures. KEY RESULTS AND CONCLUSION: The concentration of digestive hairs along vascular bundles in subgenus Genlisea is a primitive feature, indicating its basal position within the genus. Digestive hairs of Genlisea consist of three compartments with different ultrastructure and function. In subgenus Tayloria the terminal hair cells are transfer cells, but not in species of subgenus Genlisea. A digestive pool of viscous fluid occurs in Genlisea traps. In spite of their similar architecture, the digestive-absorptive hairs of Lentibulariaceae feature differences in morphology and ultrastructure.     

Eimeria trichosuri: Phylogenetic position of a marsupial coccidium, based on 18S rDNA sequences

Phylogenetic analysis of the genus Eimeria suggests that parasite and host have coevolved over broad evolutionary timescales. Here we extend this analysis by determining the 18S rDNA gene sequence of the marsupial coccidium, Eimeria trichosuri, and assessing its phylogenetic position relative to Eimeria from birds, reptiles and placental mammals. This analysis placed E. trichosuri clones in a clade that diverged before the major clade comprising species from placental mammals. The position of E.trichosuri is consistent with host phylogeny where marsupials represent an ancient evolutionary line that predates the placental mammal line.     

Floral micromorphology and nectar composition of the early evolutionary lineage Utricularia (subgenus Polypompholyx,Lentibulariaceae)

Protoplasma - Utricularia (Lentibulariaceae) is a genus comprising around 240 species of herbaceous, carnivorous plants. Utricularia is usually viewed as an insect-pollinated genus, with the...     

Floral micromorphology of the Australian carnivorous bladderwort <Emphasis Type="Italic">Utricularia dunlopii</Emphasis>, a putative pseudocopulatory species

Flowers of sexually deceptive taxa generally possess a set of morphological and physiological characters that mimic their insect pollinators. These characters often include a specific insect-like floral configuration, together with scent glands (osmophores) that produce fragrances which chemically resemble insect sex pheromones. Furthermore, these flowers tend not to produce pollinator food rewards. According to some authors, flowers of the Australian bladderwort Utricularia dunlopii (and species of the Utricularia capilliflora complex) resemble insects, and pollination perhaps occurs by pseudocopulation. The aims of this paper are to compare the structure and distribution of floral glandular trichomes in the Australian carnivorous plant U. dunlopii with those of closely related species assigned to the same section and to discuss their putative function. Floral tissues of U. dunlopii P. Taylor, Utricularia paulinae Lowrie, Utricularia dichotoma Labill. and Utricularia uniflora R.Br. (section Pleiochasia) were investigated using light microscopy, scanning electron microscopy, transmission electron microscopy and histochemistry. In U. dunlopii, two long, erect, filiform appendages arising from the upper lip of the corolla, together with three arising from the lower lip, bear numerous glandular trichomes that may function as osmophores. In other species, such as U. uniflora and U. paulinae, glandular papillae on the corolla palate may also function as osmophores. The floral anatomical and morphological organisation of U. dunlopii differs from that of the other investigated species, indicating that its insect pollinators are also likely to differ. Morphological and ultrastructural observations, while generally contributing to our understanding of the flower of U. dunlopii, do not refute the possibility that pollination here may occur by pseudocopulation. Further field-based investigations, however, are now necessary to test this hypothesis.     

Toward understanding the different function of two types of parenchyma cells in bamboo culms

The bamboo, woody monocot, has two types of parenchyma cells in the ground tissues of its culm, in contrast to a single type of parenchyma cell in rice, maize and other major crop species. The distribution of cell wall components, including lignin, (1-->3), (1-->4)-beta-D-glucans (MGs), the highly-substituted glucuronoarabinoxylans (hsGAXs) and low-branched xylans (lbXs) in ground parenchyma tissue of Phyllostachys heterocycla var. pubescens culms was studied at various developmental stages using light microscopy (LM), UV-microscopy, transmission electron microscopy (TEM) and immunolabeling techniques. The short parenchyma cell walls were lignified in 2-month-old bamboo culms just as the long parenchyma cell walls were. The lignified regions were confined to the portions in contact with the long parenchyma cell walls, while the walls at the cell corner region never lignified, even in 7-year-old culms. Significant differences were also found in the hemicellulose distribution between the short and long parenchyma cell walls. In bamboo parenchyma tissue, MGs were localized in short parenchyma cell walls and few were found in long parenchyma cell walls in both young and 7-year-old culms. The distribution of hsGAXs was similar to that of MGs in young culms, but they only appeared in the cell corner region of short parenchyma cells in old culms. Low-branched xylans were distributed in the lignified, but not in unlignified parenchyma cell walls. Based on this evidence, the differences of function in both short and long parenchyma cells in a bamboo culm are discussed.     

Growth and development of larvae and galls of Urophora cardui (Diptera,Tephritidae) on Cirsium arvense (Compositae)

Summary The tephritid fly Urophora cardui induces a large multi-chambered gall within the stems of Cirsium arvense. Three distinct phases of gall development have been identified as initiation, growth, and maturation. During initiation the insect gains control of tissue development and during the gall's growth phase parenchyma cells proliferate rapidly surrounding the larvae with thick layers of cells. Patches of primary nutritive cells appear along the surface of larval chambers during the growth phase but few of these cells are consumed. In the gall's maturation phase, thick layers of secondary nutritive cells appear around the surface of larval chambers and the remaining gall parenchyma lignifies. Secondary nutritive cells are the primary food of U. cardui.The gall expands rapidly during the growth phase then abruptly slows at the beginning of the maturation phase. Rate of gall growth is dependent upon the number of larvae per gall but the number of larvae does not affect duration of this phase.Larvae remain in the second instar throughout the growth phase and grow slowly. Once the gall enters the maturation phase and the secondary nutritive cells appear, the larvae moult to the third instar and grow quickly. Larvace attain over 98% of their final weight during the maturation phase and consume all secondary nutritive cells.It is postulated that larvae do not feed extensively on primary nutritive cells since these cells play a key role in gall morphogenesis. The appearance of secondary nutritive cells stimulates larval feeding at a time when gall growth and development is finished.     

Localization of Phloem Oxidases

Among oxidases, cytochrome oxidase has been localized in mitochondria of all phloem cells, catalase has been visualized in parenchyma peroxisomes and peroxidase has been localized in cell walls and in several cell organelles. In angiosperms, peroxidase is present in all phloem cell walls; it is sensitive to cyanide inhibition excepted in sieve areas and around plasmodesmata between sieve tubes and companion cells. In some species, this cyanide resistant oxidasic activity can be localized without exogenous H₂O₂. Peroxidase is localized on ribosomes, inside vacuoles, on the tonoplast and often on the plasmalemma in companion cells and differentiating sieve elements. In young sieve cells some dictyosomes can exhibit a strong peroxidasic activity. In mature parenchyma cells peroxidase can be associated with ER cisternae but not with vacuoles.     

Redifferentiation of leaflet tissues during midrib gall development in Copaifera langsdorffii (Fabaceae)

The tree Copaifera langsdorffii is a superhost for galling herbivores. This plant species has great morphogenetic potential, and responds differently to the stimuli of more than 20 gall-inducing insects. Among these, an undescribed species of Cecidomyiidae induces a midrib gall in which a radial cecidogenetic field is generated and the leaflet tissues redifferentiate. Our objectives were to assess the amplitude of this cecidogenetic field, in which the leaflet tissues were influenced by the feeding action of the cecidomyiid; how the final gall shape was generated; and if tissue redifferentiation conferred any adaptive value on the galling herbivore. Leaflet morphogenesis followed the pattern described in the literature for simple leaves, resulting in a mesophytic arrangement. Tissue redifferentiation due to gall formation revealed that in a midrib gall, abaxial epidermal cells divided to enlarge the gall; spongy parenchyma cells originated the storage tissue, secretory structures, and vascular bundles; palisade parenchyma cells became homogeneous; and adaxial epidermis originated the nutritive tissue. Cell elongation, a necessary step towards cell redifferentiation, is triggered by an increase in water transport to the gall site and vacuole pressure due to neoformed xylem bundles. The generation of the final shape of the midrib gall involved repetitive histological steps in response to the amplitude of the cecidogenetic field. The largest impact of the cecidomyiid feeding action occurred in gall tissues redifferentiated from protoderm and adaxial ground meristem, which provided advantages to the gall maker of the C. langsdorffii midrib gall in terms of nutritional value, microenvironment, and protection against natural enemies.     

Microsatellite markers developed for Utricularia reniformis (Lentibulariaceae)

? Premise of the study: Microsatellite markers were developed to study the genetic diversity and population structure of the carnivorous bladderwort Utricularia reniformis, which is endemic to the Atlantic Forest of southern and southeastern Brazil. Cross-species amplification was tested in U. gibba, U. neottioides, U. subulata, and Pinguicula benedicta. ? Methods and Results: The U. reniformis genome was sequenced in a 454 GS FLX sequencer, and eight primer sets were developed based on the microsatellites identified from the reads. All loci are polymorphic, showing 1.6 to 4.8 alleles per population. Preliminary results show that primer sets are suitable for population-level studies. Cross-species amplification was successful in three other Utricularia species and one Pinguicula species. ? Conclusions: Markers developed in this study provide tools for analyses of intra- and interpopulation genetic diversity in Utricularia and Pinguicula.