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Main Conclusion: The anatomical structures of Carex moorcroftii roots showing stronger plasticity during drought had a lower coefficient of variation in cell size in the same habitats, while those showing weaker plasticity had a higher coefficient of variation. The complementary relationship between these factors comprises the adaptation mechanism of the C. moorcroftii root to drought. To explore the effects of habitat drought on root anatomy of hygrophytic plants, this study focused on roots of C. moorcroftii. Five sample plots were set up along a soil moisture gradient in the Western Sichuan Plateau to collect experimental materials. Paraffin sectioning was used to obtain root anatomy, and one-way ANOVA, correlation analysis, linear regression analysis, and RDA ranking were applied to analyze the relationship between root anatomy and soil water content. The results showed that the root transverse section area, thickness of epidermal cells, exodermis and Casparian strips, and area of aerenchyma were significantly and positively correlated with soil moisture content (P < 0.01). The diameter of the vascular cylinder and the number and total area of vessels were significantly and negatively correlated with the soil moisture content (P < 0.01). The plasticity of the anatomical structures was strong for the diameter and area of the vascular cylinder and thickness of the Casparian strip and epidermis, while it was weak for vessel diameter and area. In addition, there was an asymmetrical relationship between the functional adaptation of root anatomical structure in different soil moisture and the variation degree of root anatomical structure in the same soil moisture. Therefore, the roots of C. moorcroftii can shorten the water transport distance from the epidermis to the vascular cylinder, increase the area of the vascular cylinder and the number of vessels, and establish a complementary relationship between the functional adaptation of root anatomical structure in different habitats and the variation degree of root anatomical structure in the same habitat to adapt to habitat drought. This study provides a scientific basis for understanding the response of plateau wetland plants to habitat changes and their ecological adaptation strategies. More scientific experimental methods should be adopted to further study the mutual coordination mechanisms of different anatomical structures during root adaptation to habitat drought for hygrophytic plants., (© 2024. The Author(s).)

Carex muskingumensis is a highly valued perennial ornamental grass cultivated worldwide. However, there is limited genetic data regarding this species. Selection of proper reference genes (RGs) for reverse transcription quantitative PCR (RT-qPCR) data normalization has become an essential step in gene expression analysis. In this study, we aimed to examine expression stability of nine candidate RGs in C. muskingumensis plants, subjected to osmotic stress, generated either by salinity or PEG treatment. The identification of genes exhibiting high expression stability was performed by four algorithms (geNorm, NormFinder, BestKeeper and deltaCt method). The results showed that the combination of two genes would be sufficient for reliable expression data normalization. ADP (ADP-ribosylation factor) and TBP (TATA-box-binding protein) were identified as the most stably expressed under salinity treatment, while eIF4A (eukaryotic initiation factor 4A) and TBP were found to show the highest stability under PEG-induced drought. A set of three genes ( ADP , eIF4A and TBP ) displayed the highest expression stability across all experimental samples tested in this study. To our best knowledge, this is the first report regarding RGs selection in C. muskingumensis . It will provide valuable starting point information for conducting further analyses in this and related species concerning their responses to water shortage and salinity stress.

Pleistocene climate fluctuations had profound influence on the biogeographical history of many biota. As large areas in high mountain ranges were covered by glaciers, biota were forced either to peripheral refugia (and possibly beyond to lowland refugia) or to interior refugia (nunataks). However, nunatak survival remains controversial as it relies solely on correlative genetic evidence. Here, we test hypotheses of glacial survival using two high alpine plant species (the insect-pollinated Pedicularis asplenifolia and wind-pollinated Carex fuliginosa) in the European Alps. Employing the iDDC (integrative Distributional, Demographic and Coalescent) approach, which couples species distribution modelling, spatial and temporal demographic simulation and Approximate Bayesian Computation, we explicitly test three hypotheses of glacial survival: (a) peripheral survival only, (b) nunatak survival only and (c) peripheral plus nunatak survival. In P. asplenifolia the peripheral plus nunatak survival hypothesis was supported by Bayes factors (BF> 100), whereas in C. fuliginosa the peripheral survival only hypothesis, although best supported, could not be unambiguously distinguished from the peripheral plus nunatak survival hypothesis (BF = 5.58). These results are consistent with current habitat preferences (P. asplenifolia extends to higher elevations) and the potential for genetic swamping (i.e., replacement of local genotypes via hybridization with immigrating genotypes [expected to be higher in the wind-pollinated C. fuliginosa]). Although the persistence of plants on nunataks during glacial periods has been debated and studied over decades, this is one of the first studies to explicitly test the hypothesis instead of solely using correlative evidence., (© 2019 The Authors. Molecular Ecology published by John Wiley & Sons Ltd.)

A Gram-stain-negative strictly aerobic, marine bacterium, designated GH2-2 T , was isolated from a rhizosphere mudflat of a halophyte (Carex scabrifolia) in Gangwha Island, the Republic of Korea. The cells of the organism were oxidase-positive, catalase-positive, flagellated, short rods that grew at 10-40°C, pH 4-10, and 0-13% (w/v) NaCl. The predominant ubiquinone was Q-10. The major polar lipids were phosphatidylcholine, phosphatidylethanolamine, and phosphatidylglycerol. The major fatty acid is C 18:1 . Phylogenetic analysis based on 16S rRNA gene sequences revealed that the novel isolate formed an independent lineage at the base of the radiation encompassing members of the genus Thioclava, except for Thioclava arenosa. The closest relatives were T. nitratireducens (96.03% sequence similarity) and T. dalianensis (95.97%). The genome size and DNA G+C content were 3.77 Mbp and 59.6 mol%, respectively. Phylogenomic analysis supported phylogenetic distinctness based on 16S rRNA gene sequences. Average nucleotide identity values were 73.6-74.0% between the novel strain and members of the genus Thioclava. On the basis of data obtained from a polyphasic approach, the strain GH2-2 T (= KCTC 62124 T = DSM 105743) represents a novel species of a new genus for which the name Hahyoungchilella caricis gen. nov., sp. nov. is proposed. Moreover, the transfer of Thioclava arenosa Thongphrom et al. 2017 to Pseudothioclava gen. nov. as Pseudothioclava arenosa comb. nov. is also proposed. Finally, Thioclava electrotropha Chang et al. 2018 is proposed to be a later heterosynonym of Thioclava sediminum Liu et al. 2017.

Heavy metal contamination, one of the greatest global problems, not only endangers humans and animals but also negatively affects plants. New trends, the production and industrial applications of metals in nanoforms, lead to release of large amounts of nanoparticles into the environment. However, the influence of nanoparticles on living organisms is not well understood. Cadmium is a heavy metal not essential for plants, and to its phytotoxicity also contributes its chemical similarity to zinc. It has been recorded that zinc at low concentrations reduces the toxicity of cadmium, but our results with ZnO nanoparticles did not proved it. In contrast, ZnO nanoparticles significantly increased the negative effect of cadmium, which was reflected mainly in changes in the content of photosynthetic pigments.

Salt stress is a major abiotic stress threatening plant growth and development throughout the world. In this study, we investigated the salt stress adaptation mechanism of Carex rigescens (Franch.) V. Krecz, a stress-tolerant turfgrass species with a wide distribution in northern China. Specifically, we analyzed the growth, physiology, and transcript expression patterns of two C. rigescens genotypes (Huanghua and Lvping No.1) exposed to salt stress. Results show that Huanghua demonstrated better growth performance, and higher turf quality (TQ), photochemical efficiency (Fv/Fm), relative water content (RWC), proline content, and lower relative electrolyte leakage (REL) during seven days of salt treatment compared to Lvping No.1, suggesting that Huanghua is more salt tolerant. Significant differences in reactive oxygen species (ROS), Malondialdehyde (MDA), melatonin, non-enzymatic antioxidants, lignin, and flavonoid content, as well as in antioxidant activity between Huanghua and Lvping No.1 after salt stress indicate the diverse regulation involved in salt stress adaptation in C. rigescens. These results, combined with those of the transcript expression pattern of involved genes, suggest that Huanghua is more active and efficient in ROS scavenging, Ca 2+ binding, and its phytohormone response than Lvping No.1. Meanwhile, Lvping No.1 showed relatively higher phenylpropanoid synthesis, using flavonoid and lignin as supplements for the inadequate ROS-scavenging capacity and the development of vascular tissues, respectively. These performances illustrate the differences between the two genotypes in multifaceted and sophisticated actions contributing to the tolerance mechanism of salt stress in C. rigescens. In addition, the significantly higher content of melatonin and the rapid induction of Caffeic acid O-methyltransferase (COMT) highlight the role of melatonin in the salt stress response in Huanghua. The results of our study expand existing knowledge of the complexity of the salt stress response involving the antioxidant system, Ca 2+ signaling, phytohormone response signaling, and phenylpropanoid pathways. It also provides a basis for further study of the underlying mechanism of salt tolerance in C. rigescens and other plant species., (Copyright © 2018 Elsevier GmbH. All rights reserved.)

Background: Forage nutritive value plays an important role in livestock nutrition and maintaining sustainable grassland ecosystems, and grazing management can affect the quality of forage. In this study, we investigated the effects of different grazing intensities on the nutritive values of Leymus chinensis (Trin.) Tzvelev, Artemisia spp. and Carex duriuscula C. A. Mey in the steppes of China during the growing seasons from 2011 to 2013. Five grazing management treatments were implemented: (1) rest grazing in spring, heavy grazing in summer and moderate grazing in autumn (RHM), (2) rest grazing in spring, moderate grazing in summer and heavy grazing in autumn (RMH), (3) heavy grazing though all seasons (HHH), (4) heavy grazing in spring and summer and moderate grazing in autumn (HHM) and (5) continuous moderate grazing in all seasons (MMM)., Results: There were significant effects of year, season, treatment, and year × season and year × treatment interactions only on the crude protein of L. chinensis (P < 0.05). The crude protein concentrations of L. chinensis in the plots of constant high grazing pressure (HHH) and reduced grazing pressure in the last grazing stage (HHM) were higher than with deferred grazing (RMH and RHM, P < 0.05) in spring from 2011 to 2012. For Artemisia spp. and C. duriuscula, the crude protein concentration in HHH was higher than that in RMH (P < 0.05) in the summer of 2011. There were no significant differences (P > 0.05) for ether extract, neutral detergent fiber, acid detergent fiber and Ca concentration for any of the grasses in spring and summer from 2011 to 2013 under the different grazing management treatments., Conclusions: The nutritive value of L. chinensis was more responsive to grazing disturbance than Artemisia spp. and C. duriuscula, and heavy grazing maintained a relatively high crude protein content in all species. Seasonal and interannual seasonal differences in grazing management combinations were two of the most important factors in determining the variability of forage nutritive value, including crude protein, ether extract, neutral detergent fiber, acid detergent fiber and calcium, for L. chinensis, Artemisia spp. and C. duriuscula. We suggest that moderate grazing should be adopted to ensure the quality and yield of forage and promote the sustainable development of animal husbandry.

Bipolar disjunct distributions are a fascinating biogeographic pattern exhibited by about 30 vascular plants, whose populations reach very high latitudes in the northern and southern hemispheres. In this review, we first propose a new framework for the definition of bipolar disjunctions and then reformulate a list of guiding principles to consider how to study bipolar species. Vicariance and convergent evolution hypotheses have been argued to explain the origin of this fragmented distribution pattern, but we show here that they can be rejected for all bipolar species, except for Carex microglochin. Instead, human introduction and dispersal (either direct or by mountain-hopping)-facilitated by standard and nonstandard vectors-are the most likely explanations for the origin of bipolar plant disjunctions. Successful establishment after dispersal is key for colonization of the disjunct areas and appear to be related to both intrinsic (e.g., self-compatibility) and extrinsic (mutualistic and antagonistic interactions) characteristics. Most studies on plant bipolar disjunctions have been conducted in Carex (Cyperaceae), the genus of vascular plants with the largest number of bipolar species. We found a predominant north-to-south direction of dispersal, with an estimated time of diversification in agreement with major cooling events during the Pliocene and Pleistocene. Bipolar Carex species do not seem to depend on specialized traits for long-distance dispersal and could have dispersed through one or multiple stochastic events, with birds as the most likely dispersal vector., (© 2017 Botanical Society of America.)

Premise of the Study: Bipolar species represent the greatest biogeographical disjunction on Earth, raising many questions about the colonization and adaptive processes behind such striking distribution. We investigated climatic niche differences of five Carex bipolar species in North and South America to assess niche shifts between these two regions. Moreover, we assessed potential distribution changes with future climate change., Methods: We used 1202 presence data points from herbarium specimens and 19 bioclimatic variables to assess climatic niche differences and potential distributions among the five species using ordination methods and Maxent., Key Results: The niche overlap analyses showed low levels of niche filling and high climatic niche expansion between North and South America. Carex macloviana and C. maritima showed the greatest niche expansion (60% and 96%, respectively), followed by C. magellanica (45%) and C. microglochin (39%). Only C. canescens did not colonize new environments (niche expansion = 0.2%). In contrast, all species but C. magellanica had niche filling that was <40%; hence, they are absent in the south from many environments they inhabit in North America. Climate change will push all species toward higher latitudes and elevation, reducing the availability of suitable environments., Conclusions: The colonization of South America seems to have involved frequent climatic niche shifts. Most species have colonized new environments from those occupied in the North. Observed niche shifts appear congruent with time since colonization and with current genetic structure within species. In these cold-dwelling species, climate change will most likely decrease their suitable environments in the future., (© 2017 Botanical Society of America.)

We examined foliar nitrogen (N) and phosphorus (P) stoichiometry of 3 wetland plants (Phalaris arundinacea, Miscanthus sacchariflorus, and Carex brevicuspis) distributed along an elevation gradient in the Dongting Lake, China, and how this stoichiometry is related to soil physico-chemical characteristics, elevation, and flooding days. Plant and soil samples were collected from 3 lakeshore sites. Total N and P concentrations of plants and six physico-chemical characteristics of the soil were measured, in addition to the elevation and flooding days. P. arundinacea and M. sacchariflorus had higher total N and P concentrations than C. brevicuspis. The foliar N:P ratio decreased with increasing elevation, and only increased with increasing foliar total N concentration. Canonical correspondence analysis indicated that the foliar stoichiometry was primarily regulated by soil water content, followed by soil nutrient concentration. The foliar N and P stoichiometry of the 3 wetland plants was insignificantly correlated with soil total P concentration. However, foliar stoichiometric characteristics and soil total N concentration significantly differed among the 3 species. These results demonstrate that spatial variation of foliar stoichiometry in wetland plants exists along an elevation gradient, with this information being useful for the conservation and management of wetland plants in this lake.

The study of migration pathways of artificial isotopes in the flood-plain biogeocoenoses, impacted by the nuclear fuel cycle plants, requires determination of isotope speciations in the biomass of higher terrestrial plants. The optimal method for their determination is the sequential elution technique (SET). The technique was originally developed to study atmospheric pollution by metals and has been applied to lichens, terrestrial and aquatic bryophytes. Due to morphological and physiological differences, it was necessary to adapt SET for new objects: coastal macrophytes growing on the banks of the Yenisei flood-plain islands in the near impact zone of Krasnoyarsk Mining and Chemical Combine (KMCC). In the first version of SET, 20 mM Na 2 EDTA was used as a reagent at the first stage; in the second version of SET, it was 1 M CH 3 COONH 4 . Four fractions were extracted. Fraction I included elements from the intercellular space and those connected with the outer side of the cell wall. Fraction II contained intracellular elements; fraction III contained elements firmly bound in the cell wall and associated structures; fraction IV contained insoluble residue. Adaptation of SET has shown that the first stage should be performed immediately after sampling. Separation of fractions III and IV can be neglected, since the output of isotopes into the IV fraction is at the level of error detection. The most adequate version of SET for terrestrial vascular plants is the version using 20 mM Na 2 EDTA at the first stage. Isotope 90 Sr is most sensitive to the technique changes. Its distribution depends strongly on both the extractant used at stage 1 and duration of the first stage. Distribution of artificial radionuclides in the biomass of terrestrial vascular plants can vary from year to year and depends significantly on the age of the plant., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Premise of the Study: Understanding the drivers of speciation is a central task of evolutionary biology. Chromosomal rearrangements are known to play an important role in species diversification, but the role of rearrangements of holocentric chromosomes-chromosomes without localized centromeres-is poorly understood., Methods: We made numerous artificial crosses between Carex scoparia individuals of different diploid chromosome numbers and, for comparison, between individuals of the same chromosome number. We studied chromosome pairing and chromosomal rearrangements in the F1 individuals using light microscopy. We then estimated germination rates as a function of geographic distance, genetic distance, chromosome number differences in parents, and pairing irregularities in F1 individuals, using generalized least squares to fit alternative regression models., Key Results: The most informative predictors of germination rates in the F1 generation are chromosome number differences and minimum number of chromosome pairing irregularities in the F1 individuals. Genetic and geographic distances between parents are not significant predictors., Conclusions: Holocentric chromosomal rearrangements play an important role in postzygotic reproductive isolation in Carex through F1 hybrid inviability and sterility. Hybrid dysfunction seems to be a suitable model for chromosomal speciation when there are several chromosomal rearrangements between parents. However, we have not tested the hypothesis that genome rearrangements may also play an important role in suppressing recombination between cytogenetically divergent populations., (© 2016 Botanical Society of America.)

Wetland creation has become a commonplace method for mitigating the loss of natural wetlands. Often mitigation projects fail to restore ecosystem services of the impacted natural wetlands. One of the key ecosystem services of newly created wetlands is carbon accumulation/sequestration, but little is known about how planting diversity (PD) affects the ability of herbaceous wetland plants to store carbon in newly created wetlands. Most mitigation projects involve a planting regime, but PD, which may be critical in establishing biologically diverse and ecologically functioning wetlands, is seldom required. Using a set of 34 mesocosms (∼1 m(2) each), we investigated the effects of planting diversity on carbon storage potential of four native wetland plant species that are commonly planted in created mitigation wetlands in Virginia - Carex vulpinoidea, Eleocharis obtusa, Juncus effusus, and Mimulus ringens. The plants were grown under the four distinctive PD treatments [i.e., monoculture (PD 1) through four different species mixture (PD 4)]. Plant biomass was harvested after two growing seasons and analyzed for tissue carbon content. Competition values (CV) were calculated to understand how the PD treatment affected the competitive ability of plants relative to their biomass production and thus carbon storage potentials. Aboveground biomass ranged from 988 g/m(2) - 1515 g/m(2), being greatest in monocultures, but only when compared to the most diverse mixture (p = 0.021). However, carbon storage potential estimates per mesocosm ranged between 344 g C/m(2) in the most diverse mesocosms (PD 4) to 610 g C/m(2) in monoculture ones with no significant difference (p = 0.089). CV of E. obtusa and C. vulpinoidea showed a declining trend when grown in the most diverse mixtures but J. effusus and M. ringens displayed no difference across the PD gradient (p = 0.910). In monocultures, both M. ringens, and J. effusus appeared to store carbon as biomass more effectively than the other species, suggesting that the choice of plant species may play an important role in facilitating the development of carbon accumulation/storage in created wetlands. Plant community diversity provides many ecosystem services (e.g., habitat and floristic quality) other than carbon storage function. Thus, a further study is needed that will focus on investigating how other design elements such as microtopography and hydrologic connectivity may interact with PD in terms of enhancing the carbon storage potential of newly created wetlands., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

The trade-off between allocation to sexual and clonal reproduction in clonal plants is influenced by a variety of environmental factors; however, it has rarely been examined under field conditions. In this study, we investigated the trade-off between two modes of reproduction in Carex brevicuspis C. B. Clarke across a small-scale elevational gradient (21-27 m a.s.l.) at the Dongting Lake wetlands, China. The proportion of biomass allocated to and the density of reproductive ramets were higher at low than at intermediate and high elevations. In contrast, the proportion of biomass allocated to and the density of rhizome buds were lower at low than at intermediate and high elevations. Redundancy analysis showed that sexual reproduction was positively correlated with soil moisture content, soil organic matter, total phosphorus, and pH, and negatively correlated with elevation and ramet density. Our findings suggested that allocation to sexual reproduction is favored in disturbed habitats with fertile soils, whereas allocation to vegetative propagation is favored in stable and competitive habitats. Trade-off between allocation to sexual reproduction and vegetative propagation along an elevational gradient might be a reproductive strategy of C. brevicuspis to adapt to the water level fluctuations in wetland habitats.

Background and Aims: Environmental temperature regulates plant regeneration via seed in several superimposed ways, and this complex regulation will be disrupted by climate change. The role of diurnally alternating temperatures (ΔT) in terminating dormancy will be a major factor in this disruption, as its effects on seed germination are immediate., Methods: The effect of ΔT on seed germination was modelled using two populations of the wetland sedge Carex diandra, one from a montane site and one from a subalpine site. A cardinal-temperature model was fitted to germination results obtained from a thermal gradient plate, and the model was used to simulate changes in germination under two possible future climate scenarios (RCP2·6 and RCP8·5, for representative concentration pathways) as defined by the Intergovernmental Panel on Climate Change., Key Results: Scenario RCP2·6 projected moderate increases in average temperatures and ΔT, whereas RCP8·5 projected greater warming and higher ΔT. Increasing ΔT decreased the base temperature for seed germination and the thermal time required for germination. The effect of higher ΔT together with the higher temperatures increased germination under both climate scenarios., Conclusions: Carex diandra germination is highly responsive to potential changes in ΔT, and thus this study highlights the role of ΔT in seed responses to climate change. Comprehensive cardinal-temperature models, encompassing the different effects of temperature on seed germination, are needed to understand how climate change will affect plant regeneration., (© The Author 2015. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Unlabelled: •, Premise of the Study: Most alpine plants in the Northeast United States are perennial and flower early in the growing season, extending their limited growing season. Concurrently, they risk the loss of reproductive efforts to late frosts. Quantifying long-term trends in northeastern alpine flower phenology and late-spring/early-summer frost risk is limited by a dearth of phenology and climate data, except for Mount Washington, New Hampshire (1916 m a.s.l.).•, Methods: Logistic phenology models for three northeastern US alpine species (Diapensia lapponica, Carex bigelowii and Vaccinium vitis-idaea) were developed from 4 yr (2008-2011) of phenology and air temperature measurements from 12 plots proximate to Mount Washington's long-term summit meteorological station. Plot-level air temperature, the logistic phenology models, and Mount Washington's climate data were used to hindcast model yearly (1935-2011) floral phenology and frost damage risk for the focal species.•, Key Results: Day of year and air growing degree-days with threshold temperatures of -4°C (D. lapponica and C. bigelowii) and -2°C (V. vitis-idaea) best predicted flowering. Modeled historic flowering dates trended significantly earlier but the 77-yr change was small (1.2-2.1 d) and did not significantly increase early-flowering risk from late-spring/early-summer frost damage.•, Conclusions: Modeled trends in phenological advancement and sensitivity for three northeastern alpine species are less pronounced compared with lower elevations in the region, and this small shift in flower timing did not increase risk of frost damage. Potential reasons for limited earlier phenological advancement at higher elevations include a slower warming trend and increased cloud exposure with elevation and/or inadequate chilling requirements., (© 2014 Botanical Society of America, Inc.)

In a seven-year study, we tested effects of increased N and O3 deposition and climatic conditions on biomass of subalpine grassland. Ozone risk was assessed as exposure (AOT40) and as stomatal flux (POD0,1). We hypothesized that productivity is higher under N- and lower under O3 deposition, with interactions with climatic conditions. Aboveground biomass was best correlated with growing-degree days for May (GDDMay). Nitrogen deposition increased biomass up to 60% in the highest treatment, and 30% in the lowest addition. Also belowground biomass showed a positive N-response. Ozone enrichment had no effect on biomass, and no interaction between O3 and N was observed. Growth response to N deposition was not correlated to GDDMay or precipitation, but indicated a cumulative effect over time. Productivity of subalpine grassland is tolerant to increasing ozone exposure, independent of N input and climatic drivers. N deposition rates at current critical loads, strongly increase the grassland yield., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

In this study, the soil moisture content was measured, and the quantitative characteristics of this sedge species were compared. The phenotypic plasticity of each parameter and the linear regression relationships were analyzed. The results showed that the soil moisture content was significantly affected by location, soil depth, and sampling date. The aboveground biomass, underground biomass, biomass density, and population density at the peak were significantly higher than elsewhere on the dune. However, the morphological plasticity index of the quantitative characteristics was higher at the base and middle of the dune. When the soil moisture content decreased, the underground biomass and ramet biomass density increased. The aboveground and underground biomasses were strongly negatively correlated, but the ramet height and aboveground biomass were strongly positively correlated. These results indicated that the soil water content significantly affected the clonal growth of C. physodes. The responsiveness of C. physodes may be adaptive when the soil resource supply is low. The strong morphological plasticity of the species appears to be ecologically important for the maintenance and dominance of this species in the dune habitat.

Background: Introduced earthworms are widespread in forests of North America creating significant negative impacts on forest understory communities. However, much of the reported evidence for negative earthworm effects comes from field investigations either comparing invaded and non-invaded forests or across invasion fronts. While important, such work is rarely able to capture the true effect of earthworms on individual plant species because most forests in North America simultaneously face multiple stressors which may confound earthworm impacts.We used a mesocosm experiment to isolate effects of the anecic introduced earthworm, Lumbricus terrestris L. on seedlings of 14 native plant species representing different life form groups (perennial herb, graminoid, and tree)., Results: Earthworm presence did not affect survival, fertility or biomass of any of the seedling plant species tested over a 17-week period. However, L. terrestris presence significantly decreased growth of two sedges (Carex retroflexa Muhl. ex Willd. and Carex radiata (Wahlenb.) Small) by decreasing the number of culms., Conclusions: Our mesocosm results with seedlings contrast with field reports indicating extensive and significant negative effects of introduced earthworms on many mature native forbs, and positive effects on sedges. We suggest that earthworm impacts are context- and age-specific and that generalizations about their impacts are potentially misleading without considering and manipulating other associated factors.

The response of Carex lasiocarpa in riparian wetlands in Sanjiang Plain to the environmental gradient of water depth was analyzed by using the Gaussian Model based on the biomass and average height data, and the ecological water-depth amplitude of Carex lasiocarpa was derived. The results indicated that the optimum ecological water-depth amplitude of Carex lasiocarpa based on biomass was [13.45 cm, 29.78 cm], while the optimum ecological water-depth amplitude of Carex lasiocarpa based on average height was [2.31 cm, 40.11 cm]. The intersection of the ecological water-depth amplitudes based on biomass and height confirmed that the optimum ecological water-depth amplitude of Carex lasiocarpa was [13.45 cm, 29.78 cm] and the optimist growing water-depth of Carex lasiocarpa was 21.4 cm. The TWINSPAN, a polythetic and divisive classification tool, was used to classify the wetland ecological series into 6 associations. Result of TWINSPAN matrix classification reflected an obvious environmental gradient in these associations: water-depth gradient. The relation of biodiversity of Carex lasiocarpa community and water depth was determined by calculating the diversity index of each association.

Plant communities often exhibit high diversity, even though pairwise experiments usually result in competitive hierarchies that should result in competitive exclusion. Such experiments, however, do not typically allow expression of spatial traits, despite theoretical studies showing the potential importance of spatial mechanisms of diversity maintenance. Here we ask whether, in a clonal plant model system, spatial trait variation is more likely than growth trait variation to maintain diversity. We used a field-calibrated, spatially explicit model to simulate communities comprising sets of four simulated species differing in only one of a suite of architectural or growth traits at a time, examining their dynamics and long-term diversity. To compare trait manipulation effects across traits measured in different units, we scaled traits to have identical effects on initial productivity. We found that in communities of species differing only in an architectural trait, all species usually persist, whereas communities of species differing only in a growth trait experienced rapid competitive exclusion. To examine the roles of equalizing and stabilizing mechanisms in maintaining diversity, we conducted reciprocal invasion experiments for species pairs differing only in single traits. The results suggest that stabilizing mechanisms cannot account for the observed long-term co-occurrence. Strong positive correlations between diversity and similarity both in monoculture carrying capacity and reciprocal invasion ability suggesting equalizing mechanisms may instead be responsible.

The experiment with seeds buried in soil has been carried out for 63 alpine plant species from the Northwest Caucasus. Seeds were mixed with native soils and placed in soil at the depth of 8-10 cm for five years. After excavation, seeds of 45 species did not germinate at all. Viability of eight species, four Carex species among them, exceeded 10%. These species are typical of Geranium-Hedysarum meadows and alpine snowbeds and form the main part of soil seed banks in these communities.

Data from four components of the radiation balance were used to investigate the surface energy budgets for a Carex lasiocarpa mire in the Sanjiang Plain, Northeast China, and the controlling factors of the evapotranspiration (ET) were discussed in detail. During the growing season 2006, the shortwave radiation (SW↓) reaching the mire surface added up to 2,854.3 MJ m(-2) and the net radiation (Rn) was 1,637.4 MJ m(-2) in total, with an average of 9.86 MJ m(-2) day(-1). G was the smallest flux at the water-atmosphere interface, with an average of about 0.91 MJ m(-2) day(-1), but showed high relative variability, even changing its sign. The latent and sensible heat fluxes (LE and H) amounted to 787.48 and 476.26 MJ m(-2), respectively, and the total sum of LE and H accounted for 77.18% of Rn. By conversion from LE, the average value of ET from the mire was 1.84 mm day(-1), amounting to 298.8 mm. The total ET was almost 60% of the total rainfall in the same period, proving that ET is the primary water consumer in the mire. The growth of C. lasiocarpa was related closely with surface resistance (r (s)), and analysis of partial correlation indicated that r (s) correlated negatively with leaf area index (LAI) when the interference of the available energy, Rn-G, was removed. There was a strong linkage between r (s) and the evaporative fraction [LE/(LE + H)] as well as Bowen ratio (β). r (s) was the key factor in controlling the variation of ET and regulating energy partitioning between LE and H. During the whole growing season, r (s) and R (n)-G were the two main factors coupled in ET processes. In spring, r (s) dominated ET processes, and the increase in LAI led to a decrease in r (s), which in turn accelerated ET as vegetation developed until late August. After August, the available energy controlled the process of ET completely until ET reached an equilibrium in mid-October.

Hemiparasites are known to influence community structure and ecosystem functioning, but the underlying mechanisms are not well studied. Variation in the impacts of hemiparasites on diversity and production could be due to the difference in the relative strength of two interacting pathways: direct negative effects of parasitism and positive effects on N availability via litter. Strong effects of parasitism should result in substantial changes in diversity and declines in productivity. Conversely, strong litter effects should result in minor changes in diversity and increased productivity. We conducted field-based surveys to determine the association of Castilleja occidentalis with diversity and productivity in the alpine tundra. To examine litter effects, we compared the decomposition of Castilleja litter with litter of four other abundant plant species, and examined the decomposition of those four species when mixed with Castilleja. Castilleja was associated with minor changes in diversity but almost a twofold increase in productivity and greater foliar N in co-occurring species. Our decomposition trials suggest litter effects are due to both the rapid N loss of Castilleja litter and the effects of mixing Castilleja litter with co-occurring species. Castilleja produces litter that accelerates decomposition in the alpine tundra, which could accelerate the slow N cycle and boost productivity. We speculate that these positive effects of litter outweigh the effects of parasitism in nutrient-poor systems with long-lived hemiparasites. Determining the relative importance of parasitism and litter effects of this functional group is crucial to understand the strong but variable roles hemiparasites play in affecting community structure and ecosystem processes.

Fens are among the most threatened habitats in Europe as their area has decreased considerably in the last centuries. For successful management and restoration conservationists need detailed knowledge about seed bank formation and seed longevity of plants, as these features are closely related to successional and vegetation dynamical processes. I analysed seed longevity and the germination characteristics of six fen plant species by seed burial experiments. Based on seed weight, seed bank was expected for long-term persistent for the light-seeded Schoenus nigricans, Carex appropinquata, C. pseudocyperus, C. davalliana and Peucedanum palustre and also that for the medium-seeded Cicuta virosa. It was proved that, the latter two species have short-term persistent seed banks, while Carex pseudocyperus has a transient seed bank, therefore these species may only have a limited role in restoration from seed banks. It was found that Schoenus nigricans, Carex appropinquata and C. davalliana have persistent seed banks, because some of their four-year-old seeds have emerged. Fresh seeds had low germination rate in all studied species and majority of seeds emerged after winter, except for Carex pseudocyperus. After the germination peak in spring, the majority of the ungerminated seeds of Schoenus nigricans, Peucedanum palustre, Carex appropinquata, C. davalliana and Cicuta virosa entered a secondary dormancy phase that was broken in autumn. I found the seasonal emergence of the latter three species highly similar.

We investigated whether plant interaction intensity in a subarctic-alpine meadow is important for determining community structure and species abundance. Using two common species as phytometers, we measured interaction intensity using a neighbor removal approach. Eight biotic and abiotic variables known to influence species abundance and community structure were measured, with regression trees used to examine how plant interactions and the biotic and abiotic variables were related to species evenness, richness, and phytometer spatial cover. A range of interactions was present, with both strong competition and facilitation present over small-scale abiotic and biotic gradients. Despite the variation in interaction intensity, it was generally unrelated to either community structure or phytometer cover. In other words, plant interactions were intense in many cases but were not important to community structure. This may be due to the prevalence of clonal species in this system and the influence of previous year's interactions on plant survival and patterns of community structure. These results also suggest how conflicting theories of the role of competition in unproductive environments may be resolved. Our findings suggest that plant interactions may be intense in reducing individual growth, while simultaneously not important in the context of community structure. Plant interactions need to be viewed and tested relative to other factors and stresses to accurately evaluate their importance in plant communities, with continued differentiation between the intensity of plant interactions and their relative importance in communities.

The composition of isolated floras has long been thought to be the result of relatively rare long-distance dispersal events. However, it has recently become apparent that the recruitment of lineages may be relatively easy and that many dispersal events from distant but suitable habitats have occurred, even at an infraspecific level. The evolution of the flora on the high mountains of Africa has been attributed to the recruitment of taxa not only from the African lowland flora or the Cape Floristic Region, but also to a large extent from other areas with temperate climates. We used the species rich, pan-temperate genera Carex, Ranunculus and Alchemilla to explore patterns in the number of recruitment events and region of origin. Molecular phylogenetic analyses, parametric bootstrapping and ancestral area optimizations under parsimony indicate that there has been a high number of colonization events of Carex and Ranunculus into Africa, but only two introductions of Alchemilla. Most of the colonization events have been derived from Holarctic ancestors. Backward dispersal out of Africa seems to be extremely rare. Thus, repeated colonization from the Northern Hemisphere in combination with in situ radiation has played an important role in the composition of the flora of African high mountains.

Monoecy and protogyny are widespread in wind-pollinated plants and have been interpreted as outcrossing mechanisms, though few studies have investigated their function. Carex, a large genus of anemophilous herbs, is predominantly monoecious and many species are protogynous. We investigated whether monoecy and protogyny limit self-pollination in seven Carex species. We conducted field experiments comparing stigmatic pollen loads and seed set between intact and emasculated stems. We tested for self-compatibility and evaluated pollen limitation of seed set by supplemental pollination. Finally, we measured outcrossing rates in open-pollinated and emasculated stems using allozyme markers. Emasculated stems captured significantly less pollen than open-pollinated stems and set less seed. Pollen deposition during the female-only phase for intact stems was only 12% of the total captured. Outcrossing rates for three species indicated high selfing (range t = 0.03-0.39). Allozyme loci in the remaining species were monomorphic also suggesting high selfing. These results demonstrate that neither monoecy nor protogyny is particularly effective at limiting self-fertilization. Selection for the avoidance of selfing is unlikely to maintain monoecy in many Carex species although protogyny may provide limited opportunities for outcrossing. We propose that geitonogamy in self-compatible wind-pollinated species with unisexual flowers may be widespread and provides reproductive assurance.

A greenhouse mesocosm experiment, representing earthworm-free North American Acer-dominated forest floor and soil conditions, was used to examine the individual and combined effects of initial invasion by three European earthworm species (Dendrobaena octaedra, Lumbricus rubellus and Lumbricus terrestris) on the forest floor and upper soil horizons, N and P availability, and the mortality and biomass of four native understory plant species (Acer saccharum, Aquilegia canadensis, Aralia racemosa, and Carex pensylvanica). All the three earthworm species combined caused larger impacts on most variables measured than any single earthworm species. These included loss of O horizon mass, decreased thickness of the O horizon and increased thickness of the A horizon, and higher availability of N and P. The latter finding differs from field reports where nutrients were less available after invasion, and probably represents an initial transient increase in nutrient supply as earthworms consume and incorporate the O horizon into the A horizon. Earthworms also increased mortality of plants and decreased total mesocosm plant biomass, but here the impact of all the three earthworm species was no greater than that of L. terrestris and/or L. rubellus alone. This study corroborates field studies that European earthworm invasions alter North American forest ecosystem processes by initiating a cascade of impacts on plant community composition and soil properties.

Momoge wetland is an internationally important wetland not only because it is a habitat for many rare bird species but also because it is an internationally important stopover for some rare global migratory bird species. However the petroleum exploitation in wetland has brought about many environmental problems. One of the most severe problems is crude oil pollution, which has caused the dying-off Carex tato and imposes great threat on survival of rare birds. This work studied the factors that caused the dying off of Carex tato. The results showed that death of Carex tato was the result narcosis toxicity of alcohols, intermediate biodegradation products of crude oil, on the root tissue, and the fragmentization of cuticle of leaves by light components of crude oil volatized from soil surface. However, the mechanism involved was much more complex and three interactive factors including crude oil pollution of soil, long-term drought and poor permeability of soil were responsible for the dying-off of Carex tato grassland. The distribution of crude oil in soil profile was characterized with high concentrations at top silty layer and the layer below root zone and low or no crude oil at root zone layer. This distribution was relative to root system characteristics and rhizospheric biodegradation. In root zone, substantive oxygen could be transported to root zone through dense root system and well developed aerenchyma. The crude oil in root zone was easily biodegraded by aerobic rhizosphere microbes. However, some toxic intermediate products, such as some alcohols, was sealed in root zone due to poor permeability of the top soil layer and the deeper soil layer and they had lethal effects on root tissue. Above ground, low molecular components of crude oil in top soil layer was easily volatized into atmosphere during long-term drought. Some of the volatized components were adsorbed onto leaves. SEM analysis showed that these components destroyed the leaves by fragmentization of cuticle of leave. This study also shows that although wetland has natural attenuation of pollutants, natural attenuation of pollutants by natural wetland should not be overemphasized. This attenuation function may lose under some conditions such as long-term drought and poor soil texture.

Developing a relationship between the normalized difference vegetation index (NDVI) and the leaf area index (LAI) is essential to describe the pattern of spatial or temporal variation in LAI that controls carbon, water, and energy exchange in many ecosystem process models. Photosynthetic vegetation (PV) properties can affect the estimation of LAI, but no models integrate the effects of multiple species. We developed four alternative NDVI-LAI models, three of which integrate PV effects: no PV effects, leaf-level effects, canopy-level effects, and effects at both levels. The models were fit to data across the natural range of variation in NDVI for a widespread High Arctic ecosystem. The weight of evidence supported the canopy-level model (Akaike weight, wr = 0.98), which includes species-specific canopy coefficients that primarily scale fractional PV cover to LAI by accounting for the area of unexposed PV. Modeling the canopy-level effects improved prediction of LAI (R2 = 0.82) over the model with no PV effect (R2 = 0.71) across the natural range of variation in NDVI but did not affect the site-level estimate of LAI. Satellite-based methods to estimate species composition, a variable in the model, will need to be developed. We expect that including the effects of PV properties in NDVI-LAI models will improve prediction of LAI where species composition varies across space or changes over time.

Background and Aims: In wetland plant communities, species-specific responses to pulses of white light and to red : far-red light ratios can vary widely and influence plant emergence from the seed bank. Carex species are the characteristic plants of sedge meadows of natural prairie wetlands in mid-continental USA but are not returning to restored wetlands. Little is known about how light affects seed germination in these species-information which is necessary to predict seed bank emergence and to develop optimal revegetation practices. The effects of light on germination in eight Carex species from prairie wetlands were investigated., Methods: Non-dormant seeds of eight Carex species were used to determine the influence of light on germination by examining: (a) the ability of Carex seeds to germinate in the dark; (b) the effect of different lengths of exposures to white light on germination; (c) whether the effect of white light can be replaced by red light; and (d) whether the germination response of Carex seeds to white or red light is photoreversible by far-red light., Key Results: Seeds of C. brevior and C. stipata germinated >25 % in continuous darkness. Germination responses after exposure to different lengths of white light varied widely across the eight species. Carex brevior required <15 min of white light for > or =50 % germination, while C. hystericina, C. comosa, C. granularis and C. vulpinoidea required > or =8 h. The effect of white light was replaced by red light in all species. The induction of germination after exposure to white or red light was reversed by far-red light in all species, except C. stipata., Conclusions: The species-specific responses to simulated field light conditions suggest that (a) the light requirements for germination contribute to the formation of persistent seed banks in these species and (b) in revegetation efforts, timing seed sowing to plant community development and avoiding cover crops will improve Carex seed germination.

Extreme temperature events are projected to increase in frequency in a future climate. As successive extremes could occur more frequently, patches of vulnerable tundra vegetation were exposed to two consecutive heat waves (HWs) of 10 d each, with a 5-d recovery period in between. Surface temperatures during the HWs were increased approximately 6 degrees C using infrared irradiation sources. In three of the four target species (Pyrola grandiflora, Polygonum viviparum and Carex bigelowii), plant conditions improved upon the first exposure. Depending on species, leaf relative growth, leaf chlorophyll content or maximal photochemical efficiency was increased. In P. grandiflora the positive effects of the heat on the photosynthetic apparatus led to augmented net photosynthesis. By contrast, Salix arctica responded mainly negatively, indicating species-specific responses. During the second HW, leaf mortality suddenly increased, indicating that the heat stress induced by the extreme events lasted too long and negatively influenced the species resistance to high temperature. After the HWs, when plants were exposed to (low) ambient temperatures again, plant performance deteriorated further, indicating possible loss of cold resistance.

Shifts from nitrogen (N)- to phosphorus (P)-limited growth due to high N deposition may alter the functioning of wetland vegetation. This experiment tested how N vs P deficiency affects the growth and nutrient use of wetland sedges. Five wetland Carex species were grown at nine N : P supply ratios (0.6-405) with two absolute levels of N and P. Biomass and nutrient concentrations were determined after one and two growing seasons. Shoot biomass was maximal at N : P supply ratios of 15-26 after one season but 5-15 after two seasons. Photosynthesis after the first season, second-year growth, leaf longevity, and the fraction of nutrient supply retained by plants over two seasons were all negatively related to N : P supply ratios, with small effects of absolute supply. The five Carex species responded similarly to N : P ratios but differed in nutrient resorption efficiency and biomass allocation. Plants treated with high N : P ratios appeared to lose nutrients below ground. Such losses may reduce plant performance in P-limited wetlands affected by high N deposition.

Background and Aims: Plants adjust the distribution of photosynthetic capacity and chlorophyll to canopy density. The importance of the gradient in the red : far-red ratio (R : FR) relative to the irradiance gradient was studied for its perception with respect to this partitioning of photosynthetic resources. Whether the relative importance of these two signals varied between six species of different growth habit (Phaseolus vulgaris, Lysimachia vulgaris, Hedera helix, Ficus benjamina, Carex acutiformis and Brachypodium pinnatum) was investigated further., Methods: Single leaves of plants were shaded in daylight by a spectrally neutral filter or a leaf. In another experiment, leaves were treated with supplemental FR. In most cases, treatment effects were evaluated after 2 weeks., Key Results: Nitrogen and photosynthetic capacity (Amax) per leaf area, parameters pertaining to between-leaf resource partitioning, were strongly reduced in neutral shade but not additionally by spectral leaf shade. Supplemental FR reduced these parameters also, except in Carex. Acceleration of induction of senescence was observed in spectral leaf shade in primary bean leaves. Amax per unit chlorophyll, a parameter pertaining to within-leaf resource partitioning, was reduced in neutral shade, but not in spectral leaf shade or supplemental FR., Conclusions: Signalling mechanisms associated with perception of the R : FR gradient in canopies were less important than those associated with the irradiance gradient for between-leaf and within-leaf partitioning of photosynthetic resources. The relative importance of the signals differed between species because Carex was the only species for which no indications were found for an involvement of the spectral gradient in perception of canopy density.

Small habitat size and spatial isolation may cause plant populations to suffer from genetic drift and inbreeding, leading to a reduced fitness of individual plants. We examined the germination, establishment, growth, and reproductive capacity of two characteristic species of mown fen meadows, Carex davalliana, and Succisa pratensis, common in Switzerland. Plants were grown from seeds, which were collected in 18 habitat islands, differing in size and in degree of isolation. We used both common garden and reciprocal transplant experiments to assess effects of habitat fragmentation. In the common garden, plants of Carex originating from small habitat islands yielded 35% less biomass, 30% fewer tillers, and 45% fewer flowering tillers than plants from larger ones. In contrast, plants of Succisa originating from small habitat islands yielded 19% more biomass, 14% more flower heads and 35% more flowers per flower head than plants from larger ones. Moreover, plants of Succisa from small isolated habitats yielded 32% more rosettes than did plants from small connected islands. Reciprocally transplanted plants of Succisa originating from small habitat islands produced 7% more rosettes than plants from larger ones. There was no effect of small habitat size and isolation on germination and establishment of both species in the field. Our results document genetic differences in performance attributable to habitat fragmentation in both species. We suggest that fitness loss in Carex is caused by inbreeding depression, whereas in Succisa the differences in fitness are more likely caused by genetic differentiation. Our study implies that habitat fragmentation affects common habitat-specific species, such as Carex and Succisa, as well as rare ones.

Arbuscular mycorrhizal (AM) and dark septate endophytic (DSE) fungal colonization of Alchemilla glomerulans, Carex vaginata, Ranunculus acris ssp. pumilus and Trollius europaeus growing in low-alpine meadows in the Finnish subarctic were studied at different times during the growing season. Fungal colonization was correlated to soil soluble phosphorus (P) concentration. The influence of flower bud removal on fungal colonization was investigated in A. glomerulans, C. vaginata and R. acris and the correlation between AM and DSE colonization was studied. The fungal colonization patterns were found to be species-specific. R. acris maintained a relatively high rate of fungal colonization throughout the summer, while the rates of colonization of T. europaeus were lower and decreased towards the end of the season. A. glomerulans had constant arbuscular and vesicular colonization throughout the summer, but hyphal and DSE colonization declined towards the end of the season. C. vaginata did not form arbuscular mycorrhiza, but was colonized by DSE fungi and hyaline septate hyphae throughout the season. The soil soluble P concentration showed some seasonal variation, but was also highly variable between the study sites. Bud removal decreased arbuscular colonization of R. acris, but no unique effects were seen in any other parameters or the other species studied. The root fungal parameters correlated with soil P in some species at some sites, but no consistent trend was found. DSE colonization was positively correlated with root vesicular and hyphal colonization in some cases. The differences in fungal colonization parameters may be related to species-specific phenologies.