Ultrastructural analysis of Rhynchospora ovules: The first record of Cyperaceae megagametophyte on transmission electron microscope.

Microsporogenesis and microgametogenesis are unusual in sedges (Cyperaceae), the third largest monocotyledonous family, as three microspores are aborted in favor of a single functional microspore. However, studies using light microscopy show that megasporogenesis and megagametogenesis occur normally. Nevertheless, the lack of ultrastructural details limits our knowledge of female gametophyte development in this family. Given the importance of morphological studies of reproductive structures, ovules and megagametophytes of Rhynchospora pubera were analyzed under transmission electron microscopy for the first time. Overall, ovules presented features similar to those described for the family, but ultrastructural details revealed an absence of a clear boundary between the egg cell and the central cell cytoplasm. Most interestingly, antipodal and nucellar cells showed several signs of vacuolar cell death, which suggest that programmed autolysis in sporogenous and gametophytic tissue is common in gametophyte development in the Cyperaceae. This may be related to the reproductive success of this family.

RAD sequencing resolves the phylogeny, taxonomy and biogeography of Trichophoreae despite a recent rapid radiation (Cyperaceae).

Trichophoreae is a nearly cosmopolitan Cyperaceae tribe that contains ~17 species displaying striking variation in size, inflorescence complexity, and perianth morphology. Although morphologically distinct, the status of its three genera (Cypringlea, Oreobolopsis and Trichophorum) is controversial because recent phylogenetic studies have suggested they might not be reciprocally monophyletic. However, previous analyses have shown conflicting topologies and consistently poor support due to an initial rapid diversification of the tribe. We analysed restriction-site associated DNA sequencing (RADseq) data from nearly all species of the clade, combined with five Sanger-based markers (matK, ndhF, rps16, ETS-1f, ITS) sampled extensively within species. This approach allowed us to resolve deep and shallow relationships within Trichophoreae for the first time, despite an anomaly zone spanning several successive short branches that produced considerable gene tree incongruence. Analyses reveal a primary phylogenetic split of the tribe into two clades roughly corresponding to an East Asian-North American disjunction that dates back to the mid-Miocene, with both clades comprised of a mixture of reduced unispicate and larger taxa with highly compound inflorescences. Morphological characters traditionally used in the circumscription of Trichophoreae genera are shown to be homoplasious. Several of these characters correlate best with climatic conditions, with the most reduced species occurring in open habitats at high latitudes and altitudes. Close relatives with highly compound inflorescences are found in temperate or subtropical forest understories. Cypringlea and Oreobolopsis are deeply nested within Trichophorum, and we merge all three genera into a more broadly circumscribed Trichophorum. We also show that Scirpus filipes is another previously unrecognized East Asian species of Trichophorum with highly compound inflorescences.

Molecular phylogenetics of the genus Costularia (Schoeneae, Cyperaceae) reveals multiple distinct evolutionary lineages.

We investigated the monophyly of Costularia (25 species), a genus of tribe Schoeneae (Cyperaceae) that illustrates a remarkable distribution pattern from southeastern Africa, over Madagascar, the Mascarenes and Seychelles, to Malesia and New Caledonia. A further species, Tetraria borneensis, has been suggested to belong to Costularia. Relationships and divergence times were inferred using an existing four marker phylogeny of Cyperaceae tribe Schoeneae expanded with newly generated sequence data mainly for Costularia s.l. species. Phylogenetic reconstruction was executed using Bayesian inference and maximum likelihood approaches. Divergence times were estimated using a relaxed molecular clock model, calibrated with fossil data. Based on our results, Tetraria borneensis is not related to the species of Costularia. Costularia s.l. is composed of four distinct evolutionary lineages. Two lineages, one including the type species, are part of the Oreobolus clade, i.e. a much reduced genus Costularia restricted to southeastern Africa, Madagascar, the Mascarenes and Seychelles, and a small endemic genus from New Caledonia for which a new genus Chamaedendron is erected based on Costularia subgenus Chamaedendron. The other two lineages are part of the Tricostularia clade, i.e. a separate single-species lineage from the Seychelles for which a new genus (Xyroschoenus) is described, and Costularia subgenus Lophoschoenus. For the latter, more research is needed to test whether they are congeneric with the species placed in the reticulate-sheathed Tetraria clade.

Phytobeds with Fimbristylis dichotoma and Ammannia baccifera for treatment of real textile effluent: An in situ treatment, anatomical studies and toxicity evaluation.

Fimbristylis dichotoma, Ammannia baccifera and their co-plantation consortium FA independently degraded Methyl Orange, simulated dye mixture and real textile effluent. Wild plants of F. dichotoma and A. baccifera with equal biomass showed 91% and 89% decolorization of Methyl Orange within 60h at a concentration of 50ppm, while 95% dye removal was achieved by consortium FA within 48h. Floating phyto-beds with co-plantation (F. dichotoma and A. baccifera) for the treatment of real textile effluent in a constructed wetland was observed to be more efficient and achieved 79%, 72%, 77%, 66% and 56% reductions in ADMI color value, COD, BOD, TDS and TSS of textile effluent, respectively. HPTLC, GC-MS, FTIR, UV-vis spectroscopy and activated oxido-reductive enzyme activities confirmed the phytotrasformation of parent dye in to new metabolites. T-RFLP analysis of rhizospheric bacteria of F. dichotoma, A. baccifera and consortium FA revealed the presence of 88, 98 and 223 genera which could have been involved in dye removal. Toxicity evaluation of products formed after phytotransformation of Methyl Orange by consortium FA on bivalves Lamellidens marginalis revealed less damage of the gills architecture when analyzed histologically. Toxicity measurement by Random Amplification of Polymorphic DNA (RAPD) technique revealed bivalve DNA banding pattern in treated Methyl Orange sample suggesting less toxic nature of phytotransformed dye products.

Neolithic cultivation of water chestnuts (Trapa L.) at Tianluoshan (7000-6300 cal BP), Zhejiang Province, China.

Water chestnuts (Trapa) are frequently recovered at Neolithic sites along the Lower Yangtze River Valley and have been important components of the diets of prehistoric people. However, little systematic research has been conducted to determine their cultural and dietary importance. Excavations at the Tianluoshan site produced large quantities of well-preserved specimens, which provide an excellent collection for studying morphological changes with time. Using modern wild and domesticated water chestnuts (n = 447) as a reference, we find Neolithic samples (n = 481) at Tianluoshan are similar in shape but smaller in size compared to the domesticated species Trapa bispinosa. In particular, the Tianluoshan water chestnuts have bigger seeds than the wild species Trapa incisa. Further, water chestnuts diachronically increased in size at the Tianluoshan site with significant differences (one-way, ANOVA) observed for length (p = 7.85E-08), height (p = 3.19E-06), thickness (p = 1.2E-13), top diameter (p = 5.04E-08) and bottom diameter (p = 1.75E-05) between layers 7 (6700-6500 cal BP) and 6 (6500-6300 cal BP). These results suggest that water chestnuts were actively selected based on size (big), shape (full fruit, two round horns, wide base, etc.) and were an important non-cereal crop to the agricultural practices at the Tianluoshan site.

Networks of plants: how to measure similarity in vegetable species.

Despite the common misconception of nearly static organisms, plants do interact continuously with the environment and with each other. It is fair to assume that during their evolution they developed particular features to overcome similar problems and to exploit possibilities from environment. In this paper we introduce various quantitative measures based on recent advancements in complex network theory that allow to measure the effective similarities of various species. By using this approach on the similarity in fruit-typology ecological traits we obtain a clear plant classification in a way similar to traditional taxonomic classification. This result is not trivial, since a similar analysis done on the basis of diaspore morphological properties do not provide any clear parameter to classify plants species. Complex network theory can then be used in order to determine which feature amongst many can be used to distinguish scope and possibly evolution of plants. Future uses of this approach range from functional classification to quantitative determination of plant communities in nature.

Occurrence and evolutionary inferences about Kranz anatomy in Cyperaceae (Poales).

Cyperaceae is an angiosperm family with the greatest diversity of species with Kranz anatomy. Four different types of Kranz anatomy (chlorocyperoid, eleocharoid, fimbristyloid and rhynchosporoid) have been described for this angiosperm family, and the occurrence and structural characteristics of these types are important to trace evolutionary hypotheses. The purpose of this study was to examine the available data on Cyperaceae Kranz anatomy, emphasizing taxonomy, geographic distribution, habitat and anatomy, to infer the potential origin of the Kranz anatomy in this family. The results showed that the four types of Kranz anatomy (associated with C4 photosynthesis) in Cyperaceae emerged numerous times in unrelated phylogenetic groups. However, the convergence of these anatomical types, except rhynchosporoid, was observed in certain groups. Thus, the diverse origin of these species might result from different environmental pressures that promote photorespiration. Greater variation in occurrence of Kranz anatomy and anatomical types was observed in Eleocharis, whose emergence of the C4 pathway was recent compared with other genera in the family, and the species of this genus are located in aquatic environments.

Effects of lead on anatomy, ultrastructure and concentration of nutrients in plants Oxycaryum cubense (Poep. & Kunth) Palla: a species with phytoremediator potential in contaminated watersheds.

Lead (Pb) has been highlighted as a major pollutant of both terrestrial and aquatic ecosystems, causing negative impacts to these environments. The concentration of Pb in plants has increased in recent decades, mainly due to anthropogenic activities. This study has as a hypothesis that the species Oxycaryum cubense (Poep. & Kunth) Palla, abundant in aquatic environments, has the potential to be used a phytoremediator. The plants were grown in a hydroponic system with Pb in increasing concentrations (0, 4, 8, 16 and 32 mg l(-1)) for 15 days. Inductively coupled mass spectrometer (ICP OES) was used to determine the concentration of mineral nutrients and lead. Optical and transmission electron microscopy were used for the analysis of cellular damage induced by lead in roots and leaves. Ultrastructural alterations were observed as disorganization of thylakoids in the chloroplast and disruption of mitochondrial membranes in cells of leaf tissues of plants subjected to increasing Pb concentrations. There was accumulation of Pb, especially in the root system, affecting the absorption and translocation of some mineral nutrients analysed. In roots, there was reduction in the thickness of the epidermis in plants treated with Pb. This species was shown to be tolerant to the Pb concentrations evaluated, compartmentalizing and accumulating Pb mainly in roots. Due to these results, it may be considered a species with phytoremediation capacity for Pb, with potential rizofiltration of this metallic element in contaminated watersheds.

Structural variations among monocot emergent and amphibious species from lakes of the semi-arid region of Bahia, Brazil.

Temporary lakes are common in the semi-arid region of the State of Bahia and form water mirrors in the rainy season. In this period, various vegetal species appear having different life forms adapted to the seasonality conditions of the rainfall regime. This work surveyed the adaptive anatomical structures of some emergent and amphibious monocot species occurring in these lakes. We studied the anatomy of roots, rhizomes, leaves and scapes of Cyperus odoratus, Oxycaryum cubense, Pycreus macrostachyos (Cyperaceae) - amphibious species; and of Echinodorus grandiflorus (Alismataceae), Eichhornia paniculata (Pontederiaceae) and Habenaria repens (Orchidaceae) - emergent species. The anatomical features of the dermal, fundamental and vascular systems confirming the tendency of the adaptive convergence of these plants to temporary lacustrine the environment include: single layered epidermal cells with a thin cuticle layer in the aerial organs; the presence of air canals in all the organs; few or no supporting tissues; and less numerous conducting elements and thinner cell walls in the xylem. The reduction of the supporting tissues, the number of stomata, which can even be absent, and the number of conducting elements and the degree of cell wall lignification in the xylem of the emergent species is more accentuated than that of the amphibious species. The pattern of distribution of aerenchyma in the roots of the studied species was considered important to distinguish between amphibious and emergent life forms.

Anatomical analysis of turgescent and semi-dry resurrection plants: the effect of sample preparation on the sample, resolution, and image quality of X-ray micro-computed tomography (µCT).

Computer tomography has been used frequently for the 3-D visualization of plant anatomical traits but sample preparation has been widely neglected. Without any preparation smaller (i.e., up to 1 × 1 cm(2) ) turgescent or semi-dry plant samples (especially leaf samples) diminish the image quality of a scan due to gradual water loss and therefore constant movement. A suitable preparation for scans of turgescent and semi-dry plant samples with a high resolution µCT (<1-5 µm) has to be very thin, heat-resistant (up to 35°C), have a low attenuation coefficient, and should not alter the water content and structure of the sample. Several agents have been tested, but only a coating with vaseline conserved the water content of a plant sample efficiently. However, water molecules and vaseline both attenuate the X-ray beam, which decreases the image quality of scans of turgescent or semi-dry plant samples. Therefore, trade-offs between the spatial resolution, sample water content, sample size, and image quality have to be considered: larger samples have to be placed further away from the X-ray tube, which leads to a lower spatial resolution; water and preparation agents attenuate the X-ray beam, causing low-quality images which may be accompanied by motion artifacts compared to a scan of a dry sample, where no preparation is necessary.

X-ray microCT imaging technique reveals corm microstructures of an arctic-boreal cotton-sedge, Eriophorum vaginatum.

X-ray computed tomography (CT), a non-destructive imaging technique, has recently been effectively applied to botanical research. In this study an X-ray microCT technique was developed to allow for anatomical study of the overwintering corms of Eriophorum vaginatum, an ecologically important sedge species in arctic tussock-tundra and boreal peatlands. Using a GE Medical MS8X-130 X-ray microCT scanner, optimal imaging parameters included scanning isolated corms at 80 k Vp and 100 microA with a 3500 ms exposure time and an isotropic voxel size of 10 microm. A Gaussian blur image filter with a blur radius (sigma) of two pixels was applied to the optimal dataset to improve visual detection and contrast of tissues while removing 99.2% of image noise. Using the developed X-ray microCT technique several undocumented anatomical characteristics of the corm were identified including the vascular connection between a parent corm and branching cormel and the 3D shape of sclereid clusters. The 3D structure of sclereid clusters was determined whereby the perimeter of their lance shape is greatly reinforced by sclereids with thicker secondary cell walls as compared to those of the interior of the cluster. The structure of sclereid clusters and their association with leaf traces suggests they may be stabilizing the corm-leaf connection to protect vascular tissues from physical damage. The proposed X-ray microCT technique is an excellent tool for determination of the 3D structure of E. vaginatum corms and may be used to detect alterations in tissue structure and chemistry in response to environmental change in this and other Cyperaceous species.

Spikelet structure and development in Cyperoideae (Cyperaceae): a monopodial general model based on ontogenetic evidence.

BACKGROUND AND AIMS: In Cyperoideae, one of the two subfamilies in Cyperaceae, unresolved homology questions about spikelets remained. This was particularly the case in taxa with distichously organized spikelets and in Cariceae, a tribe with complex compound inflorescences comprising male (co)florescences and deciduous female single-flowered lateral spikelets. Using ontogenetic techniques, a wide range of taxa were investigated, including some controversial ones, in order to find morphological arguments to understand the nature of the spikelet in Cyperoideae. This paper presents a review of both new ontogenetic data and current knowledge, discussing a cyperoid, general, monopodial spikelet model. METHODS: Scanning electron microscopy and light microscopy were used to examine spikelets of 106 species from 33 cyperoid genera. RESULTS: Ontogenetic data presented allow a consistent cyperoid spikelet model to be defined. Scanning and light microscopic images in controversial taxa such as Schoenus nigricans, Cariceae and Cypereae are interpreted accordingly. CONCLUSIONS: Spikelets in all species studied consist of an indeterminate rachilla, and one to many spirally to distichously arranged glumes, each subtending a flower or empty. Lateral spikelets are subtended by a bract and have a spikelet prophyll. In distichously organized spikelets, combined concaulescence of the flowers and epicaulescence (a newly defined metatopic displacement) of the glumes has caused interpretational controversy in the past. In Cariceae, the male (co)florescences are terminal spikelets. Female single-flowered spikelets are positioned proximally on the rachis. To explain both this and the secondary spikelets in some Cypereae, the existence of an ontogenetic switch determining the development of a primordium into flower, or lateral axis is postulated.

Can differences in phosphorus uptake kinetics explain the distribution of cattail and sawgrass in the Florida Everglades?

BACKGROUND: Cattail (Typha domingensis) has been spreading in phosphorus (P) enriched areas of the oligotrophic Florida Everglades at the expense of sawgrass (Cladium mariscus spp. jamaicense). Abundant evidence in the literature explains how the opportunistic features of Typha might lead to a complete dominance in P-enriched areas. Less clear is how Typha can grow and acquire P at extremely low P levels, which prevail in the unimpacted areas of the Everglades. RESULTS: Apparent P uptake kinetics were measured for intact plants of Cladium and Typha acclimated to low and high P at two levels of oxygen in hydroponic culture. The saturated rate of P uptake was higher in Typha than in Cladium and higher in low-P acclimated plants than in high-P acclimated plants. The affinity for P uptake was two-fold higher in Typha than in Cladium, and two- to three-fold higher for low-P acclimated plants compared to high-P acclimated plants. As Cladium had a greater proportion of its biomass allocated to roots, the overall uptake capacity of the two species at high P did not differ. At low P availability, Typha increased biomass allocation to roots more than Cladium. Both species also adjusted their P uptake kinetics, but Typha more so than Cladium. The adjustment of the P uptake system and increased biomass allocation to roots resulted in a five-fold higher uptake per plant for Cladium and a ten-fold higher uptake for Typha. CONCLUSIONS: Both Cladium and Typha adjust P uptake kinetics in relation to plant demand when P availability is high. When P concentrations are low, however, Typha adjusts P uptake kinetics and also increases allocation to roots more so than Cladium, thereby improving both efficiency and capacity of P uptake. Cladium has less need to adjust P uptake kinetics because it is already efficient at acquiring P from peat soils (e.g., through secretion of phosphatases, symbiosis with arbuscular mycorrhizal fungi, nutrient conservation growth traits). Thus, although Cladium and Typha have qualitatively similar strategies to improve P-uptake efficiency and capacity under low P-conditions, Typha shows a quantitatively greater response, possibly due to a lesser expression of these mechanisms than Cladium. This difference between the two species helps to explain why an opportunistic species such as Typha is able to grow side by side with Cladium in the P-deficient Everglades.

Rhizome phyllosphere oxygenation in Phragmites and other species in relation to redox potential, convective gas flow, submergence and aeration pathways.

Underground rhizomes of emergent aquatic macrophytes are important for perennation, vegetative spread, competition and anchorage. In four species we examined the potential for the development of oxidized phyllospheres around rhizome apical buds, similar to the protective oxygenated rhizospheres around roots. Redox potentials and polarographic measurements of radial oxygen loss were recorded using platinum cathodes around the apical buds. The aeration pathway from atmosphere to phyllosphere was investigated anatomically and by applied pressurized gas flow. Redox potentials increased by +400, +45, +200 and +340 mV around rhizome apices of Phragmites australis, Oryza rhizomatis, Carex rostrata and Glyceria maxima, respectively. Radial oxygen loss from rhizome apices of Phragmites was increased by convective gas flow through the rhizome and by shoot de-submergence, and decreased by resistances applied within the aeration pathway and by shoot submergence. We conclude that oxygen passes via internal gas-space connections between aerial shoot, rhizome and underground buds and into the phyllosphere regions via scale-leaf stomata and surfaces on the buds. We suggest that oxidized phyllospheres may protect rhizome apices against phytotoxins in waterlogged soils, just as oxidized rhizospheres protect roots.

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

When grown in nutrient solutions of extremely low [P] (

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.

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.

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.

The unusual vascular structure of the corm of Eriophorum vaginatum: implications for efficient retranslocation of nutrients.

Eriophorum spp. are abundant perennial graminoids in the Arctic tundra and boreal peatlands. Because ecological studies indicated that some plants are unusually productive on infertile and cold sites, the anatomy of the overwintering corms of Eriophorum vaginatum (L.) and Eriophorum scheuchzeri (Hoppe) were examined to determine their involvement in nutrient uptake and storage. Components of the long-distance transport pathways were identified within the plants by using histochemical techniques and transport of apoplastic and symplastic dyes. E. scheuchzeri produced a rhizome that consisted mainly of storage parenchyma cells within which collateral vascular bundles were centrally located and arranged in a circle. By contrast, E. vaginatum developed a ring of horizontally arranged xylem and phloem, in addition to axial amphivasal vascular bundles leading to the leaves, all of which were bordered by transfer cells. As shown by the transport of fluorescein in the phloem and Safranine O in the xylem, each axial bundle and adventitious root contacted the horizontal ring of vascular tissues so that solutes from one vascular bundle were translocated into the vascular ring and circulated to another vascular bundle and/or to the roots. In addition, special groups of sclereids that functioned in both phloem and xylem transport were found at the base of the leaf traces and within junctions of senescing roots. These sclereids were named 'vascular sclerenchyma' and it was hypothesized that they provide a moving end for the vascular system because the corm dies progressively from the distal end as it grows upward from the apical meristem. It was concluded that this unusual vascular system of E. vaginatum is efficient in recycling nutrients internally, which may account for its competitive advantage in infertile and cold sites.