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Ontogenetic derivation and cell differentiation in photosynthetic tissues of C3 and C4 Cyperaceae.

Four variants of Kranz anatomy occur in the Cyperaceae. Three of these anatomical types (fimbristyloid, chlorocyperoid, and eleocharoid) are unique among taxa with C(4) photosynthesis in that the photosynthetic carbon reduction tissue (PCR, functional equivalent of bundle sheath) is located within the vascular strand and is separated from the primary carbon assimilation tissue (PCA, positional equivalent of mesophyll) by the mestome sheath layer. In the fourth anatomical type, rhynchosporoid, PCR tissue is located in the position of the mestome sheath. In this study, we compared two aspects of development of PCR and PCA tissues in representatives of the C(3) and C(4) types: (1) ontogenetic derivation and (2) cellular differentiation. Analysis of the planes of cell division associated with procambial strand formation indicated that PCR tissue is always derived from the procambium, while PCA tissue is derived from the ground meristem. These cell lineages remain distinct after the initial organization of vascular strands. Analysis of cell differentiation using accumulation of cell-type-specific photosynthetic enzymes as markers of differentiation indicated that, with one exception, a low level of non-cell-specific enzyme accumulation preceded abundant and cell-specific accumulation of photosynthetic enzymes at the distal end of the leaf elongation zone. Enzyme accumulation coincided spatially (and temporally) with structural aspects of cell differentiation. Previous cladistic analyses have indicated that these anatomical types represent separate evolutionary origins of the C(4) pathway, and the differences in developmental pathways observed here reflect these independent origins from C(3) ancestors.

Journal Article↗

Steroids and triterpenes from Eleocharis acutangula and E. sellowiana (Cyperaceae).

From the hexane extract of the underground parts of Eleocharis acutangula (Roxb.) Schult., lup-20(29)-ene-3beta,16beta-diol and a mixture of campesterol, stigmasterol and sitosterol were isolated. The hexane extracts of aerial and underground parts of E. sellowiana Kunth furnished two new substances, namely neohop-13(18)-en-3alpha-ol and stigmast-22-en-3beta,6beta,9alpha-triol, together with a mixture of steroids, betulinic acid, stigmast-4-en-6beta-ol-3-one and fern-9(11)-en-3alpha-ol. The molecular structures were determined by spectral analysis (1D- and 2D-NMR experiments and MS) and comparison with literature data.

Cyperaceae↗

A phylogeny of the schoenoid sedges (Cyperaceae: Schoeneae) based on plastid DNA sequences, with special reference to the genera found in Africa.

Despite its large size (about 700 species), the australy-centred sedge tribe Schoeneae has received little explicit phylogenetic study, especially using molecular data. As a result, generic relationships are poorly understood, and even the monophyly of the tribe is open to question. In this study, plastid DNA sequences (rbcL, trnL-trnF, and rps16) drawn from a broad array of Schoeneae are analysed using Bayesian and parsimony-based approaches to infer a framework phylogeny for the tribe. Both analytical methods broadly support the monophyly of Schoeneae, Bayesian methods doing so with good support. Within the schoenoid clade, there is strong support for a series of monophyletic generic groupings whose interrelationships are unclear. These lineages form a large polytomy at the base of Schoeneae that may be indicative of past radiation, probably following the fragmentation of Gondwana. Most of these lineages contain both African and non-African members, suggesting a history of intercontinental dispersal. The results of this study clearly identify the relationships of the African-endemic schoenoid genera and demonstrate that the African-Australasian genus Tetraria, like Costularia, is polyphyletic. This pattern is morphologically consistent and suggests that these genera require realignment.

Africa↗

Genetic diversity in Cymophyllus fraserianus (Cyperaceae), a rare monotypic genus.

Cymophyllus fraserianus (Fraser's sedge) is a rare perennial evergreen herb found in late-successional forest communities in the Appalachian Mountains of the U.S. Genetic diversity was assessed at 19 allozyme loci for 12 populations sampled from the southern and central portions of the range of this primitive monotypic genus. Species variation was comparable to that found for other narrowly distributed species, although mean population genetic diversity was somewhat higher. Sixteen of the 19 loci (84%) were polymorphic, with a mean of 46% polymorphic within populations. On average, 2.7 alleles were detected per polymorphic locus. Genetic diversity within populations (Hep) averaged 0.113, whereas species' genetic diversity (Hes) was 0.131. Although a twofold range of within-population genetic diversity (Hep = 0.075-0.158) was found, less than 13% of the genetic variation was distributed among populations (G(ST) = 0.129). An indirect estimate of historical levels of gene flow (Nm = 1.69) was consistent with the high mean genetic identity (mean I = 0.96) found between populations. Despite the relictual nature of C. fraserianus, its rarity and fragmented distribution, genetic diversity within the genus is typical of herbs with similar life-history characteristics. No association was found between geographic and genetic distances between populations (r = -0.01; p > 0.9), suggesting that present day populations are relatively isolated.

Cyperaceae↗

Phylogenetic relatedness limits co-occurrence at fine spatial scales: evidence from the schoenoid sedges (Cyperaceae: Schoeneae) of the Cape Floristic Region, South Africa.

Species co-occurrence at fine spatial scales is expected to be nonrandom with respect to phylogeny because of the joint effects of evolutionary (trait convergence and conservatism) and ecological (competitive exclusion and habitat filtering) processes. We use data from 11 existing vegetation surveys to test whether co-occurrence in schoenoid sedge assemblages in the Cape Floristic Region shows significant phylogenetic structuring and to examine whether this changes with the phylogenetic scale of the analysis. We provide evidence for phylogenetic overdispersion in an alliance of closely related species (the reticulate-sheathed Tetraria clade) using both quantile regression analysis and a comparison between the mean observed and expected phylogenetic distances between co-occurring species. Similar patterns are not evident when the analyses are performed at a broader phylogenetic scale. Examination of six functional traits suggests a general pattern of trait conservatism within the reticulate-sheathed Tetraria clade, suggesting a potential role for interspecific competition in structuring co-occurrence within this group. We suggest that phylogenetic overdispersion of communities may be common throughout many of the Cape lineages, since interspecific interactions are likely intensified in lineages with large numbers of species restricted to a small geographic area, and we discuss the potential implications for patterns of diversity in the Cape.

Cyperaceae↗

Floral ontogeny in ficinia and isolepis (cyperaceae), with focus on the nature and origin of the gynophore.

BACKGROUND AND AIMS: The generic delimitations of Ficinia and Isolepis, sister genera in the Cypereae, are blurred. Typical Ficinia flowers have a lobed gynophore, which envelops the base of the nutlet, whereas in Isolepis the character is considered to be absent. Some former species of Isolepis, lacking the gynophore, were recently included in Ficinia. The floral ontogeny of representative taxa in Ficinia and Isolepis were investigated with the aim of evaluating the origin and nature of the gynophore in the Cypereae. METHODS: The spikelet and floral ontogeny in inflorescences collected in the field was investigated using scanning electron microscopy (SEM) and light microscopy (LM). KEY RESULTS: SEM images of Isolepis setacea and I. antarctica, Ficinia brevifolia, F. minutiflora, F. zeyheri and F. gracilis, and LM sections of F. radiata, show that the gynoecium in Ficinia is elevated above the flower receptacle by the development of a hypogynous stalk. From its apex, a (often three-)lobed cup is formed, which envelopes the basal part of the later nutlet. In developing flowers of I. antarctica, a rudimentary hypogynous stalk appears. In I. setacea, rudiments of a hypogynous stalk can be observed at maturity. In F. radiata and F. zeyheri, intralocular hairs are present in the micropylar zone. At the surface of developing gynoecia in flowers of F. gracilis, star-shaped cuticular structures appear which disappear again at maturity. CONCLUSIONS: The overall floral ontogeny of all species studied occurs following a typical scirpoid pattern, though no perianth primordia are formed. The gynophore in Ficinia originates as a hypogynous stalk, from which the typical gynophore lobes develop. The gynophore is not homologous with the perianth.

Cyperaceae↗

Functional significance of dauciform roots: exudation of carboxylates and acid phosphatase under phosphorus deficiency in Caustis blakei (Cyperaceae).

Caustis blakei produces an intriguing morphological adaptation by inducing dauciform roots in response to phosphorus (P) deficiency. We tested the hypothesis that these hairy, swollen lateral roots play a similar role to cluster roots in the exudation of organic chelators and ectoenzymes known to aid the chemical mobilization of sparingly available soil nutrients, such as P. Dauciform-root development and exudate composition (carboxylates and acid phosphatase activity) were analysed in C. blakei plants grown in nutrient solution under P-starved conditions. The distribution of dauciform roots in the field was determined in relation to soil profile depth and matrix. The percentage of dauciform roots of the entire root mass was greatest at the lowest P concentration ([P]) in solution, and was suppressed with increasing solution [P], while in the field dauciform roots were predominantely located in the upper soil horizons, and decreased with increasing soil depth. Citrate was the major carboxylate released in an exudative burst from mature dauciform roots, which also produced elevated levels of acid phosphatase activity. Malonate was the dominant internal carboxylate present, with the highest concentration in young dauciform roots. The high concentration of carboxylates and phosphatases released from dauciform roots, combined with their prolific distribution in the organic surface layer of nutrient-impoverished soils, provides an ecophysiological advantage for enhancing nutrient acquisition.

Acid Phosphatase↗

AFLP markers and cytotaxonomic analysis reveal hybridisation in the genus Schoenus (Cyperaceae).

Molecular, cytological, and morphological data support the existence of a hybrid population between Schoenus nigricans and Schoenus ferrugineus. This population was found in northeastern Italy, where S. nigricans is central with respect to its natural range and S. ferrugineus is marginal, being most common in the Alps and in central and northern Europe. Molecular marker data show that the putative hybrid population is genetically intermediate between nearby populations of the parent species. Cytological evidence confirmed the hybrid nature of this population, as does the almost complete sterility of plants within the population. Although no seeds were produced by the hybrid population, some possibly fertile pollen grains were produced; this suggests that the possibility of introgression between the two species through the hybrids cannot completely be excluded.

Crosses, Genetic↗

Molecular phylogeny of Japanese Eleocharis (Cyperaceae) based on ITS sequence data, and chromosomal evolution.

ITS sequence data were used to estimate the phylogeny of 24 Japanese Eleocharis species and to make karyomorphological observations on 19 of these taxa. Two major clades were identified in Japanese Eleocharis molecular phylogenetic trees: (1) one including all species of section Limnochloa, and (2) another comprising two sections, Pauciflorae and Eleocharis. Phylogenetic analysis including both Japanese and North American species also shows strong support for monophyly of the Mutatae/ Limnochloa clade. The width of the spikelets in the species Mutatae/ Limnochloa is the same as that of the culms, indicating that the relative widths of spikelets and culms are useful characteristics for classification. Two major clades were supported by karyomorphological data. All taxa of section Limnochloa had very small chromosomes, while sections Pauciflorae and Eleocharis had large chromosomes. The basic chromosome number of sections Eleocharis and Pauciflorae is thought to be x=5. Chromosomal evolution in the genus Eleocharis with diffuse centromeric chromosomes may be caused by both aneuploidization and polyploidization. Our data suggest that a 3-bp insertion near the 3' end of the 5.8S gene is useful for intrageneric delimitations of the genus Eleocharis.

Base Sequence↗

Identification and ecdysteroid antagonist activity of three oligostilbenes from the seeds of Carex pendula (Cyperaceae).

Methanolic extracts of seeds of several (Carex species were found to antagonise the action of 20-hydroxyecdysone in the Drosophila melanogaster microplate-based B(II) cell bioassay. Bioassay-guided HPLC analysis of seeds of Carex pendula (drooping sedge) provided one previously unknown tetrastilbene (cis-miyabenol A) and two known oligostilbenes (kobophenol B and cis-miyabenol C) as the biologically active compounds (EC50 values were 31, 37 and 19 microM, respectively, vs. 5 x 10(-8) M 20-hydroxyecdysone). The structures and relative stereochemistries of these compounds were deduced by comprehensive ID- and 2D-NMR experiments. These compounds are isolated from Carex pendula for the first time. In vitro experiments with dipteran and lepidopteran ecdysteroid receptor proteins demonstrate that the oligostilbenes are able to compete with radiolabelled ecdysteroid ([3H]ponasterone A) for occupancy of the ligand binding site. IC50/Ki values are similar to the EC50 values obtained in the B(II) bioassay.

Animals↗

Anticonvulsant properties of the methanolic extract of Cyperus articulatus (Cyperaceae).

The methanolic extract of rhizomes of Cyperus articulatus, a plant used in traditional medicine in Africa and Latin America for many diseases, possesses anticonvulsant activity in mice. This extract protected mice against maximal electroshock (MES)- and pentylenetetrazol (PTZ)-induced seizures. It also delayed the onset of seizures induced by isonicotinic acid hydrazide and strongly antagonized N-methyl-D-aspartate-induced turning behavior. The ED(50) for protection against seizures was 306 (154-541) mg/kg intraperitoneally (i.p.) for the PTZ test and 1005 (797-1200) mg/kg i.p. for the MES test. The ED(50) of methanolic extract against N-methyl-D-aspartate-induced turning behavior was 875 (623-1123) mg/kg i.p. C. articulatus L. methanolic extract protected 54% of mice from seizures induced by strychnine at the dose of 1000 mg/kg i.p. but had no or a moderate effect only against picrotoxin- or bicuculline-induced seizures. With these effects, the rhizome of C. articulatus L. possesses anticonvulsant properties in animals that might explain its use as a traditional medicine for epilepsy in Africa.

Animals↗

The phylogenetic position of Carex section Phyllostachys and its implications for phylogeny and subgeneric circumscription in Carex (Cyperaceae).

Wide speculation surrounds the origin and phylogenetic relationships of the most highly reduced sections in the genus Carex. In order to gain a better understanding of phylogeny in Carex, the relationship of the reduced sect. Phyllostachys to 12 putatively related sections, representing all four subgenera (Primocarex, Indocarex, Carex, Vignea), was inferred from sequences of the ITS (internal transcribed spacer) region of nrDNA. Phylogenetic reconstructions identified two main clades: (1) a "compound" clade composed of sections from subg. Indocarex, Primocarex, and a portion of subg. Carex, and (2) a "reduced" clade consisting of sections from subg. Carex (Phyllostachys) and Primocarex (Filifoliae and Firmiculmes). Subgenus Indocarex was paraphyletic within the "compound" clade supporting classifications that have merged it within a wider subg. Indocarex/Carex/Primocarex line. Subgenus Primocarex was polyphyletic. This result was consistent with theories that extreme reduction has occurred along several different evolutionary lines in Carex. Phylogenetic theories inferred from the presence or abnormal growth of the rachilla were not supported by tree topologies. Difficult sectional circumscriptions, such as the separation of sections Laxiflorae and Careyanae, were strongly upheld by sequence data. The ITS region is an effective tool for defining sectional limits and for estimating relationships among sections in Carex, but does not provide enough phylogenetic information to fully resolve relationships below the sectional level.

Journal Article↗

Fate of oxygen losses from Typha domingensis (Typhaceae) and Cladium jamaicense (Cyperaceae) and consequences for root metabolism.

The objective of this work was to determine whether radial oxygen loss (ROL) from roots of Typha domingensis and Cladium jamaicense creates an internal oxygen deficiency or, conversely, indicates adequate internal aeration and leakage of excess oxygen to the rhizosphere. Methylene blue in agar was used to visualize the pattern of ROL from roots, and oxidation of a titanium-citrate solution was used to quantify rates of oxygen leakage. Typha's roots had a higher porosity than Cladium's and responded to flooding treatment by increasing cortical air space, particularly near the root tips. A greater oxygen release, which occurred along the subapical root axis, and an increase in rhizosphere redox potential (E(h)) over time were associated with the well-developed aerenchyma system in Typha. Typha roots, regardless of oxygen release pattern, showed low or undetectable alcohol dehydrogenase (ADH) activity or ethanol concentrations, indicating that ROL did not cause internal deficiencies. Cladium roots also released oxygen, but this loss primarily occurred at the root tips and was accompanied by increased root ADH activity and ethanol concentrations. These results support the hypothesis that oxygen release by Cladium is accompanied by internal deficiencies of oxygen sufficient to stimulate alcoholic fermentation and helps explain Cladium's lesser flood tolerance in comparison with Typha.

Journal Article↗

Environmental distribution of four Carex species (Cyperaceae) in an old-growth forest.

We conducted an in-depth characterization of the range of micro-environments (1 m) in which four Carex species (C. backii, C. communis, C. plantaginea, and C. platyphylla) grow in the understory of an old-growth, deciduous forest in southern Québec, Canada. All four species occurred in significantly different micro-environments. Carex plantaginea was found at the wet end of a moisture gradient, in soils with high nitrate availability. Carex backii and C. platyphylla were found at the dry end of the moisture gradient, with C. backii occupying soils with higher phosphorus availability than C. platyphylla. Carex communis, the only ant-dispersed species studied, was found in the broadest range of environmental conditions. Our results suggest that environmental heterogeneity and interspecific microhabitat preferences are important for the maintenance of local species diversity in the forest understory, not only for common species as demonstrated in previous studies, but for infrequent species, and those within a functional group (upland Carex species). However, there was some evidence that the distributions of C. backii and C. communis were not in equilibrium with current environmental conditions, indicating that historical factors, such as dispersal and colonization events, may also have important effects on the distributions of these species and the maintenance of species diversity in old-growth forest.

Journal Article↗

Germination and establishment of forest sedges (Carex, Cyperaceae): tests for home-site advantage and effects of leaf litter.

We investigated aspects of germination and seedling establishment that might influence the distribution and diversity of Carex species growing in the forest understory. We tested the ability of Carex communis and C. platyphylla to germinate and establish at plots where adult individuals of one of these two species, or one of two other Carex species (C. backii and C. plantaginea), occurred in an old-growth forest in southern Québec, Canada. We also tested for effects of leaf litter on germination and establishment of these sedges. From a series of experiments in the field and in a lath house, we found no evidence of home-site advantage with respect to germination or seedling establishment. Leaf litter had a negative effect on germination and establishment. The results emphasize the importance of dispersal and colonization events in determining local diversity and distribution of Carex species in upland forests. High frequency of occurrence of C. communis at our study site may result from relatively wide dispersal provided by ants, and a suite of traits associated with ant dispersal in some understory Carex species.

Journal Article↗