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The floral scales in Hellmuthia (Cyperaceae, Cyperoideae) and Paramapania (Cyperaceae, Mapanioideae): an ontogenetic study.

BACKGROUND AND AIMS: In 1976 the monotypic genus Hellmuthia was placed in the Hypolytreae s.l., but was subsequently ascribed to the Mapanioideae, tribe Chrysitricheae, mainly because of the presence in Hellmuthia of two lateral, mapanioid-like floral scales with ciliated keels, the anatomy of the nutlet, the embryo and the inflorescence. Recently, based on cladistic analyses and supported by pollen ontogenetic evidence, Hellmuthia was transferred to a Cyperaceae, tribe Cypereae, clade mainly consisting of Ficinia and Isolepis. In this study, the floral ontogeny in Hellmuthia was investigated and compared with the floral ontogeny in Paramapania, with special attention for the floral scales. METHODS: Freshly collected inflorescences of Hellmuthia membranacea and Paramapania parvibractea were investigated using scanning electron and light microscopy. KEY RESULTS: In the conical 'spikelet' in Hellmuthia, proximal bracts occur, each axillating an axis with empty glume-like structures, or a reduced spikelet. Hence, it is a reduced partial inflorescence. In Hellmuthia, the stamen primordia originate before the primordia of the perianth-gynoecium appear. Moreover, a third adaxially positioned 'floral scale' was observed for the first time. The position and relative time of appearance of the floral scales in Hellmuthia are typical for perianth parts in Cyperoideae. The basal position of Hellmuthia within a clade of species with usually perianthless flowers, allows the presence of rudiments of a perianth in Hellmuthia to be interpreted as a primitive state. Development of the lateral 'scales' in Paramapania follows a different pattern. Therefore, it was decided that the lateral 'scales' in Paramapania are different from the lateral perianth parts in Hellmuthia. The pollen grains in Hellmuthia are cyperoid, with one polar and five lateral apertures, of which the membrane is covered with sexinous bodies. The pollen surface is granulate and perforate with microspines. CONCLUSIONS: The floral ontogeny in Hellmuthia occurs according to the general cyperoid pattern. The lateral scales in Hellmuthia are perianth parts, and they are not homologous to the lateral 'scales' in Paramapania.

Cyperaceae↗

The occurrence of dauciform roots amongst Western Australian reeds, rushes and sedges, and the impact of phosphorus supply on dauciform-root development in Schoenus unispiculatus (Cyperaceae).

* The incidence of species that develop specialised 'dauciform' lateral roots, which are hypothesised to be important for phosphorus (P) acquisition, is uncertain. We investigated their occurrence in Australian reed, rush and sedge species, grown at low P concentration in nutrient solution, and studied the response of Schoenus unispiculatus (Cyperaceae) to a range of P concentrations. * We assessed the fraction of root biomass invested in dauciform roots, their respiration and net P-uptake rate, and the P status of roots and leaves. * Dauciform-root development occurred only in particular genera of Cyperaceae when grown at low P supply. Increased P supply was associated with increased growth of S. unispiculatus and increased leaf [P]. Dauciform-root growth was reduced by increased P supply, and reduced P uptake co-occurred with the complete suppression of dauciform roots. * The P-induced suppression of dauciform roots in Cyperaceae is similar to that observed for proteoid roots in members of Proteaceae and Lupinus albus. The response of dauciform roots to altered P supply and their absence from root systems of some sedge species are discussed in terms of managed and natural systems.

Cyperaceae↗

The cytoskeleton and polarization during pollen development in Carex blanda (Cyperaceae).

Patterns of cytoskeletal organization during distinct polarizations that characterize pollen development in the sedge Carex blanda (Cyperaceae) were studied by correlated methods of immunohistochemistry and confocal and transmission electron microscopy. As is typical of the family Cyperaceae, Carex produces a unique pollen type known as a pseudomonad in which all four microspores of the tetrad are enclosed within the wall of a single pollen grain. Only one member of the tetrad is functional and the other three abort. The pseudomonads are precisely oriented in the locule with the functional microspore in the wide abaxial portion of the wedge-shaped cytoplasm adjacent to the tapetum, and the degenerative microspores are packed tightly in the pointed adaxial portion. A unique sequence of post-meiotic developmental events reflects both intracellular and intercellular polarity. Development includes: (1) random placement of tetrad nuclei in the coenocytic sporocyte after meiosis, (2) interrupted cytokinesis resulting in a tetrad of nuclei that migrates as a unit into the narrow adaxial tip, (3) completion of unequal cytokinesis and centering of the functional nucleus in the wide abaxial portion of the microsporocyte via a radial array of microtubules and microfilaments, (4) unequal mitosis resulting in a small generative cell at the proximal surface of the functional microspore (adjacent to the abortive microspores), and (5) recentering of the vegetative nucleus in the abaxial cytoplasm via a radial cytoskeletal array.

Journal Article↗

Mitotic microtubule development and histone H3 phosphorylation in the holocentric chromosomes of Rhynchospora tenuis (Cyperaceae).

In the present work we report the phosphorylation pattern of histone H3 and the development of microtubular structures using immunostaining techniques, in mitosis of Rhynchospora tenuis (2n = 4), a Cyperaceae with holocentric chromosomes. The main features of the holocentric chromosomes of R. tenuis coincide with those of other species namely: the absence of primary constriction in prometaphase and metaphase, and the parallel separation of sister chromatids at anaphase. Additionaly, we observed a highly conserved chromosome positioning at anaphase and early telophase sister nuclei. Four microtubule arrangements were distinguished during the root tip cell cycle. Interphase cells showed a cortical microtubule arrangement that progressively forms the characteristic pre-prophase band. At prometaphase the microtubules were homogeneously distributed around the nuclear envelope. Metaphase cells displayed the spindle arrangement with kinetochore microtubules attached throughout the entire chromosome extension. At anaphase kinetochoric microtubules become progressively shorter, whereas bundles of interzonal microtubules became increasingly broader and denser. At late telophase the microtubules were observed equatorially extended beyond the sister nuclei and reaching the cell wall. Immunolabelling with an antibody against phosphorylated histone H3 revealed the four chromosomes labelled throughout their entire extension at metaphase and anaphase. Apparently, the holocentric chromosomes of R. tenuis function as an extended centromeric region both in terms of cohesion and H3 phosphorylation.

Anaphase↗

Stilbenoids of Kobresia nepalensis (Cyperaceae) exhibiting DNA topoisomerase II inhibition.

Resveratrol oligomers, nepalensinol A, B and C, were isolated from the stem of Kobresia nepalensis (Cyperaceae). The structures were established on the basis of chemical properties and spectroscopic evidence including 2D NMR spectroscopic analysis. Nepalensinol A, B and C showed a potent inhibitory effect on topoisomerase II -- stronger than etoposide (VP-16), a topoisomerase II inhibitor used as an anti-cancer drug. Nepalensinol B, in particular, exhibited the most potent activity with an IC(50) of 0.02 microg/ml.

Animals↗

Phylogenetic relationships of Carpha and its relatives (Schoeneae, Cyperaceae) inferred from chloroplast trnL intron and trnL-trnF intergenic spacer sequences.

Within the tribe Schoeneae (Cyperaceae), the relationships between Carpha and its relatives have not been certain, and the limits and definition of Carpha have been controversial. Further, the relationships of species within Carpha have been unclear. In this study, cladistic analyses based on chloroplast trnL intron and trnL-trnF intergenic spacer sequence data were undertaken to estimate phylogenetic relationships in and around Carpha. This study found that Trianoptiles is sister to Carpha; Ptilothrix is sister to Cyathochaeta rather than to Carpha as suggested by some former authors; and Gymnoschoenus is distant from Carpha and its close relatives. The merging of Schoenoides back into Oreobolus is supported. The findings also revealed the non-monophyletic status of Costularia and of Schoenus, and indicated the phylogenetic relationships of species within Carpha.

Cyperaceae↗

Phylogenetic analysis of the nuclear alcohol dehydrogenase (Adh) gene family in Carex section Acrocystis (Cyperaceae) and combined analyses of Adh and nuclear ribosomal ITS and ETS sequences for inferring species relationships.

We analyzed sequence variation for the alcohol dehydrogenase (Adh) gene family in Carex section Acrocystis (Cyperaceae) to reconstruct Adh gene trees for Acrocystis species and to characterize the structure of the Adh gene family in Carex. Two Adh loci were included with ITS and ETS sequences in a combined Bayesian inference analysis of Carex section Acrocystis to gain a better understanding of species relationships in the section. In addition, we comment on how the results presented here contribute to our knowledge of the birth-death process of the Adh gene family in angiosperms. It appears that the structure of the Adh gene family in Carex is complex with possibly six loci present in the gene family. Additionally, variation among Acrocystis species within loci is quite low, and there is little phylogenetic resolution in the individual datasets. Bayesian inference analysis of the combined ITS, ETS, Adh1, and Adh2 datasets resulted in a moderately well-supported phylogenetic hypothesis of relationships in the section which is discussed in relation to previous hypotheses of relationships.

Alcohol Dehydrogenase↗

Flower and spikelet morphology in sawgrass, Cladium jamaicense Crantz (Cyperaceae).

In recent systematic treatments of the Cyperaceae, spikelets of all but the most primitive tribes have been considered to be indeterminate, whereas historically the number of flowers, floral sex and distribution of sexes in spikelets have been important characters in suprageneric classifications. However, descriptions of these spikelet characteristics for sawgrass, Cladium jamaicense Crantz, vary among authors. Spikelet morphology was analysed using developmental and phenological studies of sawgrass populations in south Florida, USA. Sawgrass spikelets have two flowers that expand successively. Flowers are fundamentally hermaphroditic and protogynous. The first flower to expand (F1) terminates the spikelet axis, whereas the second flower (F2), ensheathed by an addorsed prophyll, develops in the axil of the last bract produced on the axis. In 86% of the spikelets examined from ramets of three populations, the gynoecium of the F1 flower aborted, so this flower was functionally male and the spikelet was protandrous. However, in 14% of spikelets from these individuals, the F1 flower was hermaphroditic and could set seed. The F2 flower was typically hermaphroditic and matured stigmas, then anthers. Thus, spikelets in C. jamaicense are determinate and have two flowers that are dichogamous both within flowers and between flowers in a spikelet; spikelet sex expression can vary among plants and populations, especially in the first flower. These data for sawgrass suggest that a re-examination of spikelet development and phenology in other genera is needed to clarify the expression of these characters in the family.

Cyperaceae↗

Floral ontogeny in Scirpus, Eriophorum and Dulichium (Cyperaceae), with special reference to the perianth.

BACKGROUND AND AIMS: Based on molecular phylogenetic analysis, it has been suggested recently that the Cyperaceae comprises only two subfamilies: the Mapanioideae and the Cyperoideae. In most flowers of the Cyperoideae, the whorl of inner stamens is reduced, resulting in tetracyclic flowers. In the more primitive (scirpoid) genera within the Cyperoideae, the perianth consists of two polysymmetric whorls, whereas the perianth parts in the more derived genera have been subject to modifications and/or reduction. Comparative studies of the many silky hairs of Eriophorum and of the eight bristles of Dulichium have given rise to much discussion about their homology. METHODS: The spikelet and floral ontogeny in freshly collected inflorescences was investigated using scanning electron microscopy. KEY RESULTS: Complete floral ontogenies are presented for Scirpus sylvaticus L., Eriophorum latifolium Hoppe and Dulichium arundinaceum (L.) Britton, with special reference to the perianth. The results in S. sylvaticus confirm the trimerous monocot-like organization of the flower. It is used as a model for floral development in Cyperoideae. In the early developmental stages, the androecium of E. latifolium is surrounded by a massive perigonial primordium, from which the many hair-like bristles originate. Consequently, the stamens develop among the hair primordia, more or less simultaneously. The hairs are arranged in whorls, which develop centripetally. The development of the perianth in D. arundinaceum starts with the formation of three initial perianth primordia opposite the stamens. Subsequently, two more abaxial bristle primordia, alternating with the stamens, originate simultaneously with the appearance of three adaxial bristle primordia in the zone where an adaxial inner perianth primordium is expected. CONCLUSIONS: The floral development in E. latifolium and D. arundinaceum can be considered as variations upon the scirpoid floral ontogenetic theme.

Cyperaceae↗

Specialized 'dauciform' roots of Cyperaceae are structurally distinct, but functionally analogous with 'cluster' roots.

When grown in nutrient solutions of extremely low [P] (<or=1.0 microm), the sedge Schoenus unispiculatus Benth. (Cyperaceae) develops dauciform roots, which are short and carrot shaped, and produce dense numbers of long root hairs. It has been suggested that dauciform roots of monocotyledonous sedges function to acquire P from nutrient-poor, P-fixing soils in a manner similar to that of cluster (proteoid) roots developed by some dicotyledonous species, but without evidence to substantiate this claim. To elucidate the ecophysiological role of dauciform roots, we assessed carboxylate exudation, internal carboxylate and P concentrations and O(2) uptake rates during dauciform root development. We showed that O(2) consumption was fastest [9 nmol O(2) g(-1) fresh mass (FM) s(-1)] and root [P] greatest (0.4 mg P g(-1) FM) when dauciform roots were young and rapidly developing. Citrate was the most abundant carboxylate in root tissues at all developmental stages, and was most concentrated (22.2 micromol citrate g(-1) FM) in young dauciform roots, decreasing by more than half in mature dauciform roots. Peak citrate-exudation rates (1.7 nmol citrate g(-1) FM s(-1)) occurred from mature dauciform roots, and were approximately an order of magnitude faster than those from roots of species without root clusters, and similar to those of mature proteoid (cluster) roots of Proteaceae. Both developing and mature dauciform roots had the capacity to acidify (but not alkalinize) the rhizosphere. Anatomical studies showed that epidermal cells in dauciform roots were greatly elongated in the transverse plane; epidermal cells of parent roots were unmodified. Although structurally distinct, the physiology of dauciform roots in sedges appears to be analogous to that of proteoid roots of Proteaceae and Fabaceae, and hence, dauciform roots would facilitate access to sorbed P and micronutrients from soils of low fertility.

Carboxylic Acids↗

Nepalensinols D-G, new resveratrol oligomers from Kobresia nepalensis (Cyperaceae) as potent inhibitors of DNA topoisomerase II.

Four new resveratrol oligomers, nepalensinols D-G, were isolated from the stem of Kobresia nepalensis (Cyperaceae). The structures were determined by detailed NMR spectral analysis. The compounds were assessed for their inhibitory activity against human topoisomerase II, a potential target of anti-tumor agents. These stilbenoids showed potent inhibitory activity against human topoisomerase II with IC50 values of 5-15 microM.

Cyperaceae↗

Ions and amino acid analysis of Cyperus articulatus L. (Cyperaceae) extracts and the effects of the latter on oocytes expressing some receptors.

Extracts from rhizomes of Cyperus articulatus L. (Cyperaceae) used in Africa and Amazonia to treat many diseases has been shown to possess sedative and anticonvulsant properties. The aim of this study is to determine the mechanism of action of Cyperus articulatus extracts. In Xenopus oocytes expressing receptors, using electrophysiological measurement, extracts of rhizomes of Cyperus articulatus (300 microg/ml) inhibited 50% of the EC(50) and EC(80) of glutamate (1.3 and 2.9 microM, respectively) induced inward current through hNMDAR1A/2A receptors. Extracts induced very small current through rGluR3 receptors. The largest current induced by the extract (30 mg/ml) represents 128% of the EC(100) of glutamate induced inward current, through rGluR3 receptors. The excess 28% current could be induced by aspartate and/or glutamate in the extracts. The effect on Xenopus oocytes expressing heteromeric GABA(B)R1b/R2 receptors and rectifying potassium channels (Kir3) is clear. A decoction and water extract of Cyperus articulatus induced a large inward current that represented 71 and 57% (respectively) of the EC(100) of gaba (30 microM) induced inward current. The water extract induced also a large current through rectifying potassium channels (Kir3). Part of the current induced through GABA(B) receptors could be related to rectifying potassium channels and GABA(B) site receptors. Cyperus articulatus extracts possessed components that could decrease excitation (NMDA receptor antagonists) and increase inhibition (GABA(B) receptor agonists) in the central nervous system.

Amino Acids↗

Extracts from rhizomes of Cyperus articulatus (Cyperaceae) displace [3H]CGP39653 and [3H]glycine binding from cortical membranes and selectively inhibit NMDA receptor-mediated neurotransmission.

The marshland plant Cyperus articulatus (Cyperaceae) is commonly used in traditional medicine in Africa and Latin America to treat a wide variety of human diseases ranging from headache to epilepsy. We tested the hypothesis that the purported anti-epileptic effect of this plant might be due to a functional inhibition of excitatory amino acid receptors. One or several component(s) contained in the extracts inhibited the binding of [3H]CGP39653 to the NMDA recognition site and of [3H]glycine to the strychnine-insensitive glycine site of the NMDA receptor complex from rat neocortex. Water extracts from rhizomes of Cyperus articulatus dose-dependently reduced spontaneous epileptiform discharges and NMDA-induced depolarizations in the rat cortical wedge preparation at concentrations at which AMPA-induced depolarizations were not affected. We conclude that the purported beneficial effects of Cyperus articulatus might at least partially be due to inhibition of NMDA-mediated neurotransmission.

2-Amino-5-phosphonovalerate↗

Multiple locations of the rDNA sites in holocentric chromosomes of Rhynchospora (Cyperaceae).

Several cytogenetic studies have shown that representatives of the family Cyperaceae have holocentric chromosomes. Despite their interesting chromosome morphology, the chromosome organization has not been studied. This paper reports on the number and distribution of 18S-5.8S-26S ribosomal RNA sites by fluorescence in situ hybridization in eight Brazilian species of Rhynchospora. The signal of the rDNA probe was always localized in the telomeric regions. A high degree of variation was observed in the number of labelled sites, ranging from 4-8 in karyotypes with 2n = 10 to 30 sites in a karyotype with 50 chromosomes. It is possible that the same mechanism involved in the multiplication of these regions in organisms with monocentric chromosomes also plays a role in the polymorphism observed in holocentric chromosomes of Rhynchospora. An interesting feature of most hybridization sites was their diffuse state observed through to early metaphase. The decondensed state probably reflects the later transcription of this region during the cell cycle.

Chromosome Mapping↗

The genome sequence of Carex sylvatica Huds., 1762 (Poales: Cyperaceae).

We present a genome assembly of Carex sylvatica (Wood-sedge; Streptophyta; Magnoliopsida; Poales; Cyperaceae). The genome sequence has a total length of 363.24 megabases. Most of the assembly (99.79%) is scaffolded into 29 chromosomal pseudomolecules. The mitochondrial sequences have lengths of 1715.64, 74.78 and 77.37 kilobases and the plastid genome assembly has a length of 215.14 kilobases. This assembly was generated as part of the Darwin Tree of Life project, which produces reference genomes for eukaryotic species found in Britain and Ireland.

Carex sylvatica↗

The genome sequence of Schoenoplectus triqueter (L.) Palla, 1888 (Poales: Cyperaceae).

We present a genome assembly of Schoenoplectus triqueter (Triangular Club-rush; Streptophyta; Magnoliopsida; Poales; Cyperaceae). The genome sequence has a total length of 580.26 megabases. Most of the assembly (98.75%) is scaffolded into 21 chromosomal pseudomolecules. Five mitochondrial sequences and the plastid genome were also assembled. Gene annotation of this assembly on Ensembl identified 31 746 protein-coding genes. This assembly was generated as part of the Darwin Tree of Life project, which produces reference genomes for eukaryotic species found in Britain and Ireland.

Poales↗

Mycorrhizal status of the genus Carex (Cyperaceae).

The Cyperaceae have generally been considered nonmycorrhizal, although recent evidence suggests that mycotrophy may be considerably more widespread among sedges than was previously realized. This study surveyed 23 species of Carex occurring in upland and wetland habitats in northeastern Illinois. Mycorrhizal infection by arbuscular fungi was found in the roots of 16 species of Carex and appears to occur in response to many factors, both environmental and phylogenetic. While some species appear to be obligately nonmycorrhizal, edaphic influences may be responsible for infection in others. In five of the seven Carex species that were nonmycorrhizal, a novel root character, the presence of bulbous-based root hairs, was identified. The taxonomically patchy distribution of the distinctive root hair trait suggests that these structures may have evolved several times within the genus. Evidence of multiple independent origins of the root hair trait lends support to the hypothesis that root hairs represent an adaptation to nonmycotrophy. Although taxonomic position does seem to be of importance in determining the mycorrhizal dependence of sedges, the pattern may be a patchwork of both mycorrhizal clades and clades that have adapted to the nonmycorrhizal state.

Journal Article↗

The effects of population size limitation on fecundity in mosaic populations of the clonal macrophyte Scirpus maritimus (Cyperaceae).

The clonal macrophyte Scirpus maritimus (Cyperaceae) propagates locally by rhizomes and reproduces sexually by achenes. The purpose of this paper was to examine whether in size-limited habitats in patchy and discrete marshes in two Mediterranean wetlands in southern France natural populations may suffer from a reduced maternal fecundity due to a deficit in outcross pollen. We first verified that S. maritimus suffers from a reduced fecundity when self-pollinated. At a site in the Camargue, mean fecundity (mean number of achenes per centimetre of spikelet) measured in 1995 and 1996 in seven and nine populations, respectively (surface area from 50 to 4500 m) increased significantly with population surface area in 1995 but not in 1996. In the second wetland at Roquehaute, which is composed of small ponds, fecundity was very low in all 12 local populations studied in 1996 (1.1 achenes per spikelet, SD = 1.2) and was not correlated with the population surface area (from 10 to 400 m). We performed a pollen supplementation experiment in five local populations at Roquehaute to determine whether this low fecundity may be due to a pollen limitation. A significant increase in fecundity after among-pond pollinations compared to within-pond pollinations indicated that local populations suffer from a deficit in outcross pollen, since each pond appears to contain one or a few number of clones (or incompatibility types). In S. maritimus, clonal spread may have a cost in terms of reduced fecundity in small habitats because each habitat is colonized by very few clones.

Journal Article↗