Comparative flooding tolerance of Typha latifolia and Phalaris arundinacea in wetland restoration: Insights from photosynthetic CO2 response curves, photobiology and biomass allocation.

Tall helophytes such as Typha latifolia and Phalaris arundinacea often rapidly colonise after rewetting of former agricultural soil and are therefore often the first plants to contribute to the soil carbon pool. In this study we carried out a mesocosm experiment where these two species grew at three different water levels relative to the soil surface (-15 cm, 0 cm, +15 cm). After eight weeks' growth, measurements of photosynthetic CO2-response curves, stomatal conductance and chlorophyll fluorescence of photosystem II were carried out to detect flooding stress. After 10 weeks' growth, the plants were harvested and biomass production, biomass allocation and specific leaf area were determined. T. latifolia had a higher and more stable photosynthetic performance across all water level treatments, which resulted in an overall higher aboveground and belowground production than P. arundinacea. In contrast, Vcmax and Jmax decreased by 41 % and 42 %, respectively from drained to flooded conditions with signs of flooding stress as impairment of the photosynthetic apparatus. Moreover, increasing water level resulted in maintenance of aboveground organs for P. arundinacea but a decrease in allocation to belowground organs. P. arundinacea did not invest in a higher specific leaf area to counter the decreased photosynthesis under flooding. From -15 cm to 0 cm water levels, P. arundinacea showed a 68 % reduction in belowground biomass, which has negative implication for carbon retention immediately after rewetting. In contrast, recolonization of T. latifolia is likely to be a suitable contributor to the soil carbon pool due to its stable physiology and high above- and belowground biomass production at all water depths, and also likely under natural water level fluctuations. We showed that even though both species are generally considered wetland plants, they are likely to support considerably different photosynthetic carbon assimilation and soil carbon sequestration rates.

Soil salinity, not plant genotype or geographical distance, shapes soil microbial community of a reed wetland at a fine scale in the Yellow River Delta.

Soil salinization is one of the most severe environmental problems restricting biodiversity maintenance and ecosystem functioning in a coastal wetland. Recent studies have well documented how salinization affects soil microbial communities along vegetation succession of coastal wetlands. However, the salinity effect is rarely assessed in the context of plant intraspecific variation. Here, we analyzed the soil bacterial and fungal communities of Phragmites australis wetland using amplicon high-throughput sequencing at a fine scale (within 1000 m) in the Yellow River Delta. Our results revealed that microbial diversity is significantly correlated to soil salinity (assessed as electrical conductivity, EC) but not to soil nutrients (N and P content) or plant intraspecific traits (leaf length, shoot height, and neutral genetic variation). Specifically, the microbial diversity tended to decrease with increased EC, and the bacterial community was more sensitive to EC change than the fungal community. The dominant bacterial phyla were Proteobacteria, Actinobacteria, and Chloroflexi, and the dominant fungal phyla were Ascomycota, Basidiomycota, and Mortierellomycota. The relative abundance of Actinobacteria was significantly negatively correlated to EC, while Proteobacteria were positively correlated to EC. In high salinity (> 1 mS/cm), the role of the stochastic processes became more important in community assembly according to habitat niche breadth estimation, neutral community model, C-score metric, and normalized stochasticity ratio. Additional common garden and microcosm experiments provided evidence that the genotype effect of P. australis on soil microbiome might only occur between lineages from different regions but not from the same region like the Yellow River Delta. Our findings provide new insights into soil microbial community assembly processes with the intraspecific variation of host plants in the wetland ecosystem and offer a scientific reference for salinity mitigation and vegetation management of coastal wetlands under future global changes.

Can allelopathy of Phragmites australis extracts aggravate the effects of salt stress on the seed germination of Suaeda salsa?

Phragmites australis is highly adaptable with high competitive ability and is widely distributed in the coastal wetland of the Yellow River Delta. However, allelopathic effects of P. australis on the growth of neighboring plants, such as Suaeda salsa, are poorly understood. In this study, germination responses of S. salsa seeds collected from two different habitats (intertidal zone and inland brackish wetland) to the extracts from different part of P. australis were compared. Potential allelopathic effects on germination percentage, germination rate, radicle length, and seedling biomass were analyzed. The germination of S. salsa was effectively inhibited by P. australis extract. Extract organ, extract concentration, and salt concentration showed different effects, the inhibitory rates were highest with belowground extract of P. australis between the four different parts. Germination percentage and germination rate were significantly decreased by the interactive effect of salt stress and extract concentration in S. salsa from a brackish wetland but not in S. salsa from the intertidal zone. The impact of different extracts of P. australis on radicle length and seedling biomass of S. salsa showed significant but inconsistent variation. The response index results showed that the higher concentration of extract solution (50 g·L-1) of P. australis had stronger inhibitory effect on the seed germination and seedling growth of S. salsa while the belowground extract had the strongest negative effect. Our results indicated that allelopathy is an important ecological adaptation mechanism for P. australis to maintain a high interspecific competitive advantage in the species' natural habitat.

Trait value and phenotypic integration contribute to the response of exotic Rhus typhina to heterogeneous nitrogen deposition: A comparison with native Rhus chinensis.

The temporal heterogeneity of nitrogen availability in soils is increasing due to agricultural deposition. We here compared the effects of gradually increasing nitrogen deposition rate and its increasing temporal heterogeneity patterns on the functional traits of seedlings of exotic species Rhus typhina and the native species Rhus chinensis. Nitrogen deposition rates of 0, 8, 20 g N m-2 year-1 and constant, single-peak, and double-peak nitrogen were added to simulate deposition rate and temporal heterogeneity. After 60 days of treatment, R. typhina seedlings had several advantageous growth trait values, such as higher total biomass production, but lower phenotypic plasticity than R. chinensis seedlings. R. typhina seedlings also had higher phenotypic integration, measured as the correlation among functional traits. The increased nitrogen deposition rate affected several traits of the two species differently. Thus, while R. chinensis seedlings allocated more biomass to leaves and less to roots with increasing N deposition, R. typhina seedlings had stable biomass allocation among all N treatments. Chlorophyll content, leaf phosphorus concentration, and water use efficiency increased, but the maximum net photosynthetic rate decreased, with N availability in R. chinensis, but not in R. typhina. Temporal heterogeneity had no significant effect on the total biomass of R. typhina and R. chinensis seedlings. Overall, the performance of R. typhina is better than that of R. chinensis seedlings under different nitrogen deposition treatments, which is due to the significantly advantageous trait values and greater phenotypic integration of R. typhina seedlings, whereas R. chinensis seedlings have higher phenotypic plasticity.

Shade and salinity responses of two dominant coastal wetland grasses: implications for light competition at the transition zone.

BACKGROUND: Coastal wetlands are threatened by the increased salinity that may result from sea level rise. Salinity stress alters species zonation patterns through changes in competitive outcome between species differing in salinity tolerance. This study therefore aimed to understand how salinity and light affect two dominant and competing coastal wetland grasses that differ in salt tolerance, height and photosynthetic metabolism. METHODS: The C4 species Spartina anglica and the C3 species Phragmites australis were grown at five salinity levels (0, 7, 14, 21 and 28 ppt) and two light fluxes (100 % and 50 % of natural daylight) in an outdoor experimental setup for 102 d with full access to nutrients. KEY RESULTS: Salinity reduced the biomass, height and shoot density of P. australis from 81.7 g dry weight (DW), 0.73 m and 37 shoots per pot at a salinity of 0 ppt to 16.8 gDW, 0.3 m and 14 shoots per pot at a salinity of 28 ppt. Biomass, height and shoot density of S. anglica did not respond or were only slightly reduced at the highest salinity of 28 ppt. High salinity also resulted in a higher tissue concentration of N and P in P. australis. Both species had low ability to acclimate to the lower light flux. Shade acclimation in S. anglica occurred via modest changes in specific leaf area, pigment content and biomass allocation. CONCLUSIONS: High salinity reduced traits important for light competition and increased the nutrient concentration in P. australis leaf and root biomass, while this was overall unaffected in S. anglica. This is likely to reduce the competitive ability of P. australis over S. anglica for light because at high salinities the former cannot effectively shade the lower-growing S. anglica. Neither species effectively acclimates to shade, which could explain why S. anglica does not occur in the understorey of P. australis at low salinities.

Transcriptome Analysis of Tetraploid and Octoploid Common Reed (Phragmites australis).

Polyploidization in plants is thought to have occurred as coping mechanism with environmental stresses. Polyploidization-driven adaptation is often achieved through interplay of gene networks involved in differentially expressed genes, which triggers the plant to evolve special phenotypic traits for survival. Phragmites australis is a cosmopolitan species with highly variable phenotypic traits and high adaptation capacity to various habitats. The species' ploidy level varies from 3x to 12x, thus it is an ideal organism to investigate the molecular evolution of polyploidy and gene regulation mediated by different numbers of chromosome copies. In this study, we used high-throughput RNAseq data as a tool, to analyze the gene expression profiles in tetraploid and octoploid P. australis. The estimated divergence time between tetraploid and octoploid P. australis was dated to the border between Pliocene and Pleistocene. This study identified 439 up- and 956 down-regulated transcripts in tetraploids compared to octoploids. Gene ontology and pathway analysis revealed that tetraploids tended to express genes responsible for reproduction and seed germination to complete the reproduction cycle early, and expressed genes related to defense against UV-B light and fungi, whereas octoploids expressed mainly genes related to thermotolerance. Most differentially expressed genes were enriched in chaperones, folding catalysts and protein processing in endoplasmic reticulum pathways. Multiple biased isoform usage of the same gene was detected in differentially expressed genes, and the ones upregulated in octoploids were related to reduced DNA methylation. Our study provides new insights into the role of polyploidization on environmental responses and potential stress tolerance in grass species.

The river shapes the genetic diversity of common reed in the Yellow River Delta via hydrochory dispersal and habitat selection.

Understanding the driving mechanisms of local genetic diversity is a fundamental challenge under the global environmental changes. Rivers provide an excellent study system to demonstrate the effects of hydrochory dispersal and habitat selection on genetic diversity of riparian flora. In this study, we focused on the genetic variation of common reed (Phragmites australis) in the Yellow River Delta, China. Firstly, samples were collected in the Yellow River Delta, its neighboring wetland and its upstream plain. The genetic variation of P. australis was investigated using two chloroplast DNA fragments and eleven nuclear microsatellites. The findings showed that the genetic variation of P. australis in the Yellow River Delta belonged to two distinct lineages (haplotype O and haplotype P), which were similar to the upstream, and to the neighboring populations, respectively. Moreover, the genetic results suggested the potential dispersal of haplotype O from upstream to downstream. Secondly, we surveyed the plant functional traits of common reed from the Yellow River Delta in the field and in the common garden. The results showed significant differences between riverine and non-riverine populations in plant functional traits (e.g. specific leaf area and leaf length), haplotype composition and genetic clustering, which implied natural selection by habitat conditions. Lastly, we re-analyzed the plant performance data from a salt manipulation experiment with different haplotypes, and the results supported that salinity is a significant selective stressor on P. australis lineages in the Yellow River Delta. Our study highlights the significance of hydrochory dispersal and habitat selection in the river effects on genetic diversity of riparian flora, and provides important information for biodiversity conservation and wetland management in the Yellow River Delta.

Is there evidence of local adaptation of Phragmites australis to water level gradients and fluctuation frequencies?

Greater differences in hydrologic conditions are expected between coastal and inland wetlands with global climate change. Local adaptation has been considered as a significant driver of intraspecific differentiation in heterogeneous habitats. The common reed Phragmites australis is a cosmopolitan wetland species with high genetic variability and adaptability. In our study, reeds collected from coastal and inland wetlands were subjected to three stable water level gradients and two fluctuation frequencies in a common garden experiment. We measured their aboveground and belowground biomass, height, density, stem diameter, leaf water potential, specific leaf area, and photosynthetic parameters. Our results showed that P. australis exhibited high tolerance to stable and fluctuating water levels up to 30 cm depth. Increased shoot elongation rate and water-use efficiency promoted the establishment of P. australis in flooding habitats. The common reeds in the high-frequency water level fluctuation had a shorter shoot height and a lower shoot density than those in the low-frequency one. The coastal populations performed better under high (30 cm) and low (0 cm) water levels than the inland populations, which preferred shallow water (15 cm). The adaptation strategies of coastal and inland reeds to fluctuation frequencies were no different. We concluded that local adaptation might occur in P. australis populations due to different water levels rather than fluctuation frequency in coastal and inland wetlands. Our findings could provide a theoretical basis on the effects of flooding on intraspecific variation of wetland plants in future environmental change scene.

Phylogenetic diversity shapes salt tolerance in Phragmites australis estuarine populations in East China.

Estuaries are dynamic and selective environments that provide frequent opportunities for the turnover of Phragmites australis populations. We studied Phragmites genetic diversity patterns in three of the major deltas of China, viz. the Yellow River, the Yangtze and the Liaohe, in relation to Phragmites global phylogeography and soil salinity. We found that two distantly related P. australis haplotypes, each with intercontinental distribution, co-occur in these deltas in China. One is European Phragmites (Haplotype O) and is related to P. japonicus; the other (Haplotype P) has its range in East Asia and Australia and is related to the Asian tropical species P. karka. The two haplotypes have differing salt tolerance, with Haplotype O in areas with the highest salinity and Haplotype P in areas with the lowest. Introgressed hybrids of Haplotype P with P. karka, and F1 hybrids with Haplotype O, have higher salt tolerance than Haplotype P. Phylogenetic diversity appears as the factor that better explains population structure and salinity tolerance in these estuaries. Future research may explain whether the two P. australis haplotypes evolved in East Asia, and East Asia is a center of Phragmites diversity, or are introduced and a threat to P. japonicus and P. karka.

Carbon assimilation through a vertical light gradient in the canopy of invasive herbs grown under different temperature regimes is determined by leaf and whole-plant architecture.

This study examined the acclimation to temperature of two globally invasive species Iris pseudacorus and Lythrum salicaria, which share the same habitat type but differ in morphology. Iris pseudacorus has long vertical leaves, allowing light penetration through the canopy, while L. salicaria has stems with small horizontal leaves, creating significant self-shading. We aimed to build a physiological understanding of how these two species respond to different growth temperatures with regard to growth and gas exchange-related traits over the canopy. Growth and gas exchange-related traits in response to low (15 °C) and high (25 °C) growth temperature regimes were compared. Plants were grown in growth chambers, and light response curves were measured with infrared gas analysers after 23-33 days at three leaf positions on each plant, following the vertical light gradient through the canopy. After 37 days of growth, above-ground biomass, photosynthetic pigments and leaf N concentration were determined. The maximum photosynthesis rate was lower in lower leaf positions but did not differ significantly between temperatures. Iris pseudacorus photosynthesis decreased with decreasing leaf position, more so than L. salicaria. This was explained by decreasing N and chlorophyll concentrations towards the leaf base in I. pseudacorus, while pigment concentrations increased towards the lower canopy in L. salicaria. Biomass, shoot height and specific leaf area increased with temperature, more so in I. pseudacorus than in L. salicaria. Light response curves revealed that L. salicaria had a higher degree of shade acclimation than I. pseudacorus, probably due to self-shading in L. salicaria. High temperature decreased C assimilation at the bottom of the canopy in L. salicaria, while C assimilation in I. pseudacorus was less affected by temperature. As vegetative growth and flowering was stimulated by temperature, the invasive potential of these species is predicted to increase under global warming.

Nutrient removal potential and biomass production by Phragmites australis and Typha latifolia on European rewetted peat and mineral soils.

Paludiculture, sustainable and climate-smart land use of formerly drained, rewetted organic soils, can produce significant biomass in peatlands whilst potentially restoring several additional wetland services. However, the site conditions that allow maximum biomass production and nutrient removal by paludiculture crops have rarely been studied. We studied the relationship between soil characteristics, including plant-available nutrients, peak biomass, stand age, harvest period, and nutrient removal potential for two important paludiculture species, Typha latifolia and Phragmites australis, on rewetted peat and mineral soils in a large-scale European survey. T. latifolia and P. australis were able to produce an aboveground peak biomass of 10-30 t dry matter ha-1 y-1 and absorbed significant amounts of carbon, nitrogen, phosphorus, and potassium in stands older than 3 years. They were able to grow in a wide range of abiotic soil conditions. Low N:P ratios (5-9) and low N content (< 2%) in T. latifolia tissue suggest N limitation, but P uptake was still surprisingly high. P. australis had higher N:P ratios (8-25) and was less responsive to nutrients, suggesting a higher nutrient use efficiency. However, both species could still produce significant biomass at lower nutrient loads and in winter, when water content was low and nutrient removal still reasonable. Based on this European wetland survey, paludiculture holds a great potential to combine peat preservation, water purification, nutrient removal, and a high biomass production. Paludicrops take up substantial amounts of nutrients, and both summer and winter harvests provide an effective way to sequester carbon in a range of high-valued biomass products and to control nutrient effluxes from rewetted sites at the landscape scale.

Inherent trait differences explain wheat cultivar responses to climate factor interactions: New insights for more robust crop modelling.

Climate change predictions foresee a combination of rising CO2 , temperature and altered precipitation. Effects of single climatic variables are well defined, but the importance of combined variables and genotypic effects is less known, although pivotal for assessing climate change impacts, for example, with crop growth models. This study provides developmental and physiological data from combined climatic factors for two distinct wheat cultivars (Paragon and Gladius), as a basis to improve predictions for climate change scenarios. The two cultivars were grown in controlled climate chambers in a fully factorial setup of atmospheric CO2 concentration, growth temperature and watering regime. The cultivars differed considerably in their developmental rate, response pattern and the parameters responsible for most of their variation. The growth of Paragon was linked to climatic effects on photosynthesis and mainly affected by temperature. Paragon was overall more negatively affected by all treatment combinations compared to Gladius. Gladius was mostly affected by watering regime. The cultivars' acclimation strategies to climate factors varied significantly. Thus, considering a single factor is an oversimplification very likely impacting the accuracy of crop growth models. Intraspecific crop variation could help understanding genotype by environment variation. Cultivars with high phenotypic plasticity may have greater resilience against climatic variability.

Probing the Response of the Amphibious Plant Butomus umbellatus to Nutrient Enrichment and Shading by Integrating Eco-Physiological With Metabolomic Analyses.

Amphibious plants, living in land-water ecotones, have to cope with challenging and continuously changing growth conditions in their habitats with respect to nutrient and light availability. They have thus evolved a variety of mechanisms to tolerate and adapt to these changes. Therefore, the study of these plants is a major area of ecophysiology and environmental ecological research. However, our understanding of their capacity for physiological adaptation and tolerance remains limited and requires systemic approaches for comprehensive analyses. To this end, in this study, we have conducted a mesocosm experiment to analyze the response of Butomus umbellatus, a common amphibious species in Denmark, to nutrient enrichment and shading. Our study follows a systematic integration of morphological (including plant height, leaf number, and biomass accumulation), ecophysiological (photosynthesis-irradiance responses, leaf pigment content, and C and N content in plant organs), and leaf metabolomic measurements using gas chromatography-mass spectrometry (39 mainly primary metabolites), based on bioinformatic methods. No studies of this type have been previously reported for this plant species. We observed that B. umbellatus responds to nutrient enrichment and light reduction through different mechanisms and were able to identify its nutrient enrichment acclimation threshold within the applied nutrient gradient. Up to that threshold, the morpho-physiological response to nutrient enrichment was profound, indicating fast-growing trends (higher growth rates and biomass accumulation), but only few parameters changed significantly from light to shade [specific leaf area (SLA); quantum yield (φ)]. Metabolomic analysis supported the morpho-physiological results regarding nutrient overloading, indicating also subtle changes due to shading not directly apparent in the other measurements. The combined profile analysis revealed leaf metabolite and morpho-physiological parameter associations. In this context, leaf lactate, currently of uncertain role in higher plants, emerged as a shading acclimation biomarker, along with SLA and φ. The study enhances both the ecophysiology methodological toolbox and our knowledge of the adaptive capacity of amphibious species. It demonstrates that the educated combination of physiological with metabolomic measurements using bioinformatic approaches is a promising approach for ecophysiology research, enabling the elucidation of discriminatory metabolic shifts to be used for early diagnosis and even prognosis of natural ecosystem responses to climate change.

Assessing nutrient responses and biomass quality for selection of appropriate paludiculture crops.

Tall wetland graminoids with rapid growth, high productivity and wide tolerance of biotic and abiotic stresses are potentially valuable bioenergy crops, especially when grown in rewetted peat soils for biomass (paludiculture). Using wetland plants as renewable bioenergy crops instead of fossil fuels has the ecological benefits of reducing greenhouse gas (GHG) emissions, improving water quality and conserving peat soils. As these potential crops will grow in peat that differs in nutrient availability, not only will their biomass productivity be affected, but also the biomass quality for bioenergy may be altered. We set up five different nutrient availability treatments in waterlogged peat soil to simulate different nutrient environments for wetland plant cultivation. Seven wetland plants suitable for paludiculture (Typha latifolia, Arundo plinii, Arundo donax and four distinct genotypes of Phragmites australis from Denmark, The Netherlands, Romania and Italy) were selected to test responses of biomass production and tissue quality to different nutrient availability. Due to their high biomass productivity, T. latifolia, A. donax, Dutch (NL) and Romanian (RO) P. australis had the greatest potential to produce bioenergy feedstock. All taxa survived when cultivated with very low nutrient availability, especially NL and RO P. australis and T. latifolia. Moreover, biomass quality was both species-specific and element-specific, affected by increasing nutrient availability. Overall, T. latifolia had the lowest tissue concentrations of S and Si as well as high concentrations of Ca, and therefore the best tissue quality for combustion both at low and high nutrient availability. These results will provide crucial information for choosing appropriate crops and managements and promote the success of culturing wetland plants as bioenergy feedstock.

Cosmopolitan Species As Models for Ecophysiological Responses to Global Change: The Common Reed Phragmites australis.

Phragmites australis is a cosmopolitan grass and often the dominant species in the ecosystems it inhabits. Due to high intraspecific diversity and phenotypic plasticity, P. australis has an extensive ecological amplitude and a great capacity to acclimate to adverse environmental conditions; it can therefore offer valuable insights into plant responses to global change. Here we review the ecology and ecophysiology of prominent P. australis lineages and their responses to multiple forms of global change. Key findings of our review are that: (1) P. australis lineages are well-adapted to regions of their phylogeographic origin and therefore respond differently to changes in climatic conditions such as temperature or atmospheric CO2; (2) each lineage consists of populations that may occur in geographically different habitats and contain multiple genotypes; (3) the phenotypic plasticity of functional and fitness-related traits of a genotype determine the responses to global change factors; (4) genotypes with high plasticity to environmental drivers may acclimate or even vastly expand their ranges, genotypes of medium plasticity must acclimate or experience range-shifts, and those with low plasticity may face local extinction; (5) responses to ancillary types of global change, like shifting levels of soil salinity, flooding, and drought, are not consistent within lineages and depend on adaptation of individual genotypes. These patterns suggest that the diverse lineages of P. australis will undergo intense selective pressure in the face of global change such that the distributions and interactions of co-occurring lineages, as well as those of genotypes within-lineages, are very likely to be altered. We propose that the strong latitudinal clines within and between P. australis lineages can be a useful tool for predicting plant responses to climate change in general and present a conceptual framework for using P. australis lineages to predict plant responses to global change and its consequences.

Nighttime stomatal conductance differs with nutrient availability in two temperate floodplain tree species.

Nighttime water flow varies between plant species and is a phenomenon for which the magnitude, purpose and consequences are widely discussed. A potential benefit of nighttime stomata opening may be increased nutrient availability during the night since transpiration affects the mass flow of soil water towards plant roots. We investigated how nitrogen (N) and phosphorus (P) fertilization, and short-term drought affected stomatal conductance of Fraxinus excelsior L. and Ulmus laevis Pallas during the day (gs) and night (gn), and how these factors affected growth for a period of 18 weeks. Both species were found to open their stomata during the night, and gn responded to nutrients and water in a different manner than gs. Under N-deficiency, F. excelsior had higher gn, especially when P was sufficient, and lower pre-dawn leaf water potential (Ψpd), supporting our assumption that nutrient limitation leads to increases in nighttime water uptake. Under P-deficiency, F. excelsior had higher relative root production and, thus, adjusted its biomass allocation under P shortage, while sufficient N but not P contributed to overall higher biomasses. In contrast, U. laevis had higher gn and lower root:shoot ratio under high nutrient (especially N) availability, whereas both sufficient N and P produced higher biomasses. Compared with well-watered trees, the drought treatment did not affect any growth parameter but it resulted in lower gn, minimum stomatal conductance and Ψpd of F. excelsior. For U. laevis, only gs during July was lower when drought-treated. In summary, the responses of gs and gn to nutrients and drought depended on the species and its nutrient uptake strategy, and also the timing of measurement during the growing season. Eutrophication of floodplain forests dominated by F. excelsior and U. laevis may, therefore, considerably change nighttime transpiration rates, leading to ecosystem-level changes in plant-water dynamics. Such changes may have more severe consequences in the future as a higher frequency of drought events is predicted under climate change.

Influence of low calcium availability on cadmium uptake and translocation in a fast-growing shrub and a metal-accumulating herb.

Calcium (Ca) and the toxic heavy metal cadmium (Cd) are antagonistic ions competing for uptake in plants when they co-occur in soil solutions, and high Ca concentrations can reduce the uptake of Cd in plants. However, less is known about the effects of low Ca bioavailability on Cd uptake and translocation in plants. We hypothesized that low Ca availability would enhance Cd uptake and translocation in Sesbania sesban, a fast-growing shrub potentially useful for Cd removal from contaminated soils, and Brassica juncea, a well-known Cd-hyperaccumulator. The two species were grown under controlled conditions for 21 days in hydroponic nutrient solutions with either 0.2 or 2 mM Ca and 0 or 50 µM Cd in a 2 × 2 factorial experimental design. Both species had a lower relative growth rate, final root length and shoot and root biomasses at 50 µM Cd compared with growth without Cd. The negative effects of Cd on both species were exacerbated at low Ca availability. Brassica juncea had higher root Cd concentrations than S. sesban, but the shoot Cd concentrations did not differ between the two species. The low Ca concentration enhanced the uptake of Cd in the roots of both species, but Cd translocation to the shoots was low, especially at 0.2 mM Ca. We conclude that the low Ca concentration enhanced the uptake of Cd into roots of S. sesban and B. juncea and increased the phytotoxicity of Cd. The translocation of Cd to the shoots of the two species was, however, lower at 0.2 mM than at 2 mM Ca, implying that Cd removal from polluted soil cannot simply be increased by adjusting ion concentrations.

Expression of major photosynthetic and salt-resistance genes in invasive reed lineages grown under elevated CO2 and temperature.

It is important to investigate the molecular causes of the variation in ecologically important traits to fully understand phenotypic responses to climate change. In the Mississippi River Delta, two distinct, sympatric invasive lineages of common reed (Phragmites australis) are known to differ in several ecophysiological characteristics and are expected to become more salt resistant due to increasing atmospheric CO2 and temperature. We investigated whether different patterns of gene expression can explain their ecophysiological differences and increased vigor under future climatic conditions. We compared the transcript abundance of photosynthetic genes of the Calvin cycle (Rubisco small subunit, RbcS; Phosphoglycerate kinase, PGK; Phosphoribulokinase, PRK), genes related with salt transport (Na(+)/H(+) antiporter, PhaNHA) and oxidative stress response genes (Manganese Superoxide dismutase, MnSOD; Glutathione peroxidase, GPX), and the total aboveground biomass production between two genotypes representing the two lineages. The two genotypes (Delta-type, Mediterranean lineage, and EU-type, Eurasian lineage) were grown under an ambient and a future climate scenario with simultaneously elevated CO2 and temperature, and under two different soil salinities (0‰ or 20‰). We found neither differences in the aboveground biomass production nor the transcript abundances of the two genotypes, but soil salinity significantly affected all the investigated parameters, often interacting with the climatic conditions. At 20‰ salinity, most genes were higher expressed in the future than in the ambient climatic conditions. Higher transcription of the genes suggests higher abundance of the protein they code for, and consequently increased photosynthate production, improved stress responses, and salt exclusion. Therefore, the higher expression of these genes most likely contributed to the significantly ameliorated salinity impact on the aboveground biomass production of both P. australis genotypes under elevated temperature and CO2. Although transcript abundances did not explain differences between the lineages, they correlated with the increased vigor of both lineages under anticipated future climatic conditions.

Increased invasive potential of non-native Phragmites australis: elevated CO2 and temperature alleviate salinity effects on photosynthesis and growth.

The prospective rise in atmospheric CO2 and temperature may change the distribution and invasive potential of a species; and intraspecific invasive lineages may respond differently to climate change. In this study, we simulated a future climate scenario with simultaneously elevated atmospheric CO2 and temperature, and investigated its interaction with soil salinity, to assess the effects of global change on the ecophysiology of two competing haplotypes of the wetland grass Phragmites australis, that are invasive in the coastal marshes of North America. The two haplotypes with the phenotypes 'EU-type' (Eurasian haplotype) and 'Delta-type' (Mediterranean haplotype), were grown at 0‰ and 20‰ soil salinity, and at ambient or elevated climatic conditions (700 ppm CO2, +5 °C) in a phytotron system. The aboveground growth of both phenotypes was highest at the elevated climatic conditions. Growth at 20‰ salinity resulted in declined aboveground growth, lower transpiration rates (E), stomata conductance (gs), specific leaf area, photosynthetic pigment concentrations, and a reduced photosynthetic performance. The negative effects of salinity were, however, significantly less severe at elevated CO2 and temperature than at the ambient climatic conditions. The Delta-type P. australis had higher shoot elongation rates than the EU-type P. australis, particularly at high salinity. The Delta-type also had higher maximum light-saturated rates of photosynthesis (Asat), maximum carboxylation rates of Rubisco (Vcmax), maximum electron transport rates (Jmax), triose phosphate utilization rates (Tp), stomata conductance (gs), as well as higher Rubisco carboxylation-limited, RuBP regeneration-limited and Tp-regeneration limited CO2 assimilation rates than the EU-type under all growth conditions. Our results suggest that the EU-type will not become dominant over the Delta-type, since the Delta-type has superior ecophysiological traits. However, the projected rise in atmospheric CO2 and temperature will alleviate the effects of salinity on both phenotypes and facilitate their expansion into more saline areas.