Influence of die-back syndrome on reproductive strategies within Phragmites australis populations

Sexual plant reproduction is a strategy that allows plant populations to increase genetic variability, and consequently to be more efficient in adapting to new environments and to overcome stress conditions. Here, we focus on the reproductive mode of Phragmites australis, an important sub-cosmopolite species that can spread both by clonal propagation and sexual reproduction. In Europe, P. australis is affected by severe decline (known as Reed Die-Back Syndrome or RDBS); this study aims to understand if in RDBS condition the reproductive strategy of P. australis was altered, based on the hypothesis that if stress occurs, plants are likely to use their energy to increase genetic variability to overcome the stress. Inflorescences in five Italian wetlands were sampled from healthy and RDBS-affected stands of P. australis. Seed production, seed viability and seed germination were evaluated, with germination monitored under two different water conditions: moist and submerged. Stress induced by RDBS appears to influence the reproductive strategy of P. australis. In RDBS-affected stands, seed production was significantly higher than healthy stands for four of five sites. Seed viability and germination were also higher in RDBS-affected stands, but these differences were lower and site dependent.     

Assessment of aquatic and terrestrial reed (Phragmites australis) stands

A survey of recent publications shows that research on Phragmites australishas oftenapplied character because of the considerableecological and economic significance of the species.The main applications are water treatment, agriculture(food production or weed control) and natureconservation. In Europe, most research on natural reedstands has been motivated by reed die-back and effortstowards protection or restoration. Reed progressionand reed control have been the main concerns in otherparts of the world, and reed progression has alsoreceived increasing attention in Europe. While reeddie-back generally affects aquatic stands, progressioncan occur at both terrestrial and aquatic sites, andit can be desired (e.g. lake shore restoration)or unwanted (e.g. in species-rich fens ormarshes). Therefore, reed stands need to be assessedindividually to decide on management aims andappropriate methods. The varying status of Phragmites australisformed the background of the`European Reed Conference' held in Zürich/Switzerlandin October 1998. The seven contributions published inthis special issue are introduced with particularreference to differences between aquatic andterrestrial reed stands and to approaches used in theirassessment.     

Effect of cellulose wastes upon the growth of Phragmites australis

Growth responses of Phragmites australis (Cav.) Trin. Ex Steud, (reed grass), a helophyte species, were examined under in vitro and greenhouse conditions in the presence of various residues from a Kraft pulp mill. Plant tolerance to solid residues (ashes, dregs, flyashes, grits, primary sludge, and brown stock rejects) was tested in vitro. Solid residues were added separately up to 30% (w/v), as well a liquid residue up to 30% (v/v), to a Murashige and Skoog (1962) sucrose-free nutrient media with (5 mg l(-1)) 6-benzylaminopurine. After 2 mo in vitro, plantlets developed well in the presence of up to 10% solid or liquid wastes, but higher concentrations of either limited growth. This effect was mainly attributed to the plant's uptake and accumulation of various elements such as sodium, iron, copper, manganese, and boron, which are common to these waste types, thus showing an efficient phytoremediation potential. When added to MS media, the concentration of these elements generally decreased in the residual media after 2 mo of culture: the initial sodium, iron, and copper content in the growth media was reduced ca. 10-fold detected; a 5-fold reduction occurred for manganese and boron. In experiments under greenhouse conditions with in vitro propagated plantlets potted in mixtures of a commercial organic soil and residues, significant differences in plant development (plant size and fresh weight increase) were observed in the presence of ashes mixed at levels of 20% and 30%, compared to the control in organic soil. For other solid wastes, plant growth was inhibited as the concentration of each waste increased, causing chlorosis and/or plant necrosis.     

Aluminium and phosphate uptake by Phragmites australis: the role of Fe,Mn and Al root plaques

Aluminium, a potentially phytotoxic metal, is an important constituent of many mine water discharges but has largely been neglected in the literature. The behaviour of this element in the rhizosphere of the wetland plant Phragmites australis was investigated in the laboratory in the presence and absence of Mn and Fe root plaques. Electron microscopy and chemical extraction techniques were utilized to determine the physico-chemical properties of the plaques and any association of Al. Both Mn and Fe plaques occurred as amorphous coatings on root surfaces with uneven distributions. Al was not adsorbed onto the surface of either plaque type but formed a separate phosphate deposit closely resembling the Fe and Mn plaques. Phosphorus was also found to be adsorbed to the surface of the Fe plaques (but not the Mn plaques). Both mechanisms were found to immobilize P at the root surface but this did not significantly reduce the concentration of P in aerial plant tissues that was sufficient to ensure adequate growth.     

Trace elements removal ability and antioxidant activity of Phragmites australis (from Algeria)

Abstract

In this study, roots, stems and leaves of the worldwide distributed macrophyte Phragmites australis (common reed) were tested as potential removal and biomonitors of trace elements contamination in sediment. In particular, the concentrations (100, 200, and 500?mg/kg) of the following elements were analyzed: Zn, Cu, Pb, and Fe. Results showed that the amount of concentrations in plant tissues is significantly (p?≤?0.01) dependent on the kind of organ and element. Trace element concentrations decreased according to the pattern of Fe^a (Roots^a > Stems^b > Leaves^b) > Zn^b (Root^a > Leaves^b > Stems^c) > Cu^c (Roots^a > Leaves^b > Stems^c) > Pb^c (Roots^a > Stems^b > Leaves^c), as well as the roots acted as the main centers of bioaccumulation for all elements studied, and stems as the transit organs for translocation from roots to leaves. The major mechanisms employed by the plant were probably phytostabilization on the basis of the calculated Biological Concentration Factor (BCF – metal concentration ratio of plant root to soil); and Translocation Factor (TF – metal concentration ratio of plants roots to above ground part). Finally, due to the low scavenger effect of the radical DPPH, we excluded the hypothesis of the use of antioxidant mechanism in the tolerance of metals.     

Die-back of Phragmites australis: influence on the distribution and rate of sediment methanogenesis

Methanogenesis was measured during the summer of 1994, in sediment coresand bulk samples from a Phragmites australis wetland in northern Jutland,Denmark. We compared sediment from healthy reed and dying-back reed, andan open lagoon resulting from die-back. Cores revealed variability withdepth and between sites, with the highest rates coinciding with layers oforganic gyttja, and negligible methane production from the underlying sandbase. Methanogenesis rates in the lagoon and die back sites were higher(up to 100–150 nmol h^-1 g^-1dry wt. sediment) than in the healthy reed (50–80 nmolh^-1 g^-1), with the highest rates being recordedfrom May to July. At these times, methanogenesis was markedly temperature-limited; samples incubated at 30 °C anon-limiting temperature, gave rates as high as 200–400nmol h^-1 g^-1 for the lagoon and die-backareas and 150 nmol h^-1 g^-1 for the healthyarea. Addition of 8 mM acetate and H₂/CO₂headspace suggested that both acetate-fermenting andCO₂-reducing bacteria were present. Acetate additions suggested some co-limitation by substrate availability, with acetate limitation occurring in the healthy site during July and in the die-back site during August. Lower rates during August, especially in the healthy area, were associated with low water levels which resulted in more oxidized sediments. The data reveal highly variable methanogenesis in the sediment which, when considered with sediment depths, indicates that sites of Phragmites die-back have significantly greater rates of anaerobic mineralization than surrounding healthy wetland, and may be intense sources of methane.     

Freshwater wetlands: fertile grounds for the invasive Phragmites australis in a climate change context

Climate change will likely affect flooding regimes, which have a large influence on the functioning of freshwater riparian wetlands. Low water levels predicted for several fluvial systems make wetlands especially vulnerable to the spread of invaders, such as the common reed (Phragmites australis), one of the most invasive species in North America. We developed a model to map the distribution of potential germination grounds of the common reed in freshwater wetlands of the St. Lawrence River (Québec, Canada) under current climate conditions and used this model to predict their future distribution under two climate change scenarios simulated for 2050. We gathered historical and recent (remote sensing) data on the distribution of common reed stands for model calibration and validation purposes, then determined the parameters controlling the species establishment by seed. A two‐dimensional model and the identified parameters were used to simulate the current (2010) and future (2050) distribution of germination grounds. Common reed stands are not widespread along the St. Lawrence River (212 ha), but our model suggests that current climate conditions are already conducive to considerable further expansion (>16,000 ha). Climate change may also exacerbate the expansion, particularly if river water levels drop, which will expose large bare areas propitious to seed germination. This phenomenon may be particularly important in one sector of the river, where existing common reed stands could increase their areas by a factor of 100, potentially creating the most extensive reedbed complex in North America. After colonizing salt and brackishwater marshes, the common reed could considerably expand into the freshwater marshes of North America which cover several million hectares. The effects of common reed expansion on biodiversity are difficult to predict, but likely to be highly deleterious given the competitiveness of the invader and the biological richness of freshwater wetlands.     

Competition among native and invasive Phragmites australis populations: An experimental test of the effects of invasion status,genome size,and ploidy level

Among the traits whose relevance for plant invasions has recently been suggested are genome size (the amount of nuclear DNA) and ploidy level. So far, research on the role of genome size in invasiveness has been mostly based on indirect evidence by comparing species with different genome sizes, but how karyological traits influence competition at the intraspecific level remains unknown. We addressed these questions in a common‐garden experiment evaluating the outcome of direct intraspecific competition among 20 populations of Phragmites australis, represented by clones collected in North America and Europe, and differing in their status (native and invasive), genome size (small and large), and ploidy levels (tetraploid, hexaploid, or octoploid). Each clone was planted in competition with one of the others in all possible combinations with three replicates in 45‐L pots. Upon harvest, the identity of 21 shoots sampled per pot was revealed by flow cytometry and DNA analysis. Differences in performance were examined using relative proportions of shoots of each clone, ratios of their aboveground biomass, and relative yield total (RYT). The performance of the clones in competition primarily depended on the clone status (native vs. invasive). Measured in terms of shoot number or aboveground biomass, the strongest signal observed was that North American native clones always lost in competition to the other two groups. In addition, North American native clones were suppressed by European natives to a similar degree as by North American invasives. North American invasive clones had the largest average shoot biomass, but only by a limited, nonsignificant difference due to genome size. There was no effect of ploidy on competition. Since the North American invaders of European origin are able to outcompete the native North American clones, we suggest that their high competitiveness acts as an important driver in the early stages of their invasion.     

Preadaptation and post‐introduction evolution facilitate the invasion of Phragmites australis in North America

Compared with non‐invasive species, invasive plant species may benefit from certain advantageous traits, for example, higher photosynthesis capacity and resource/energy‐use efficiency. These traits can be preadapted prior to introduction, but can also be acquired through evolution following introduction to the new range. Disentangling the origins of these advantageous traits is a fundamental and emerging question in invasion ecology. We conducted a multiple comparative experiment under identical environmental condition with the invasive haplotype M lineage of the wetland grass Phragmites australis and compared the ecophysiological traits of this invasive haplotype M in North America with those of the European ancestor and the conspecific North American native haplotype E lineage, P. australis ssp. americanus. The invasive haplotype M differed significantly from the native North American conspecific haplotype E in several ecophysiological and morphological traits, and the European haplotype M had a more efficient photosynthetic apparatus than the native North American P. australis ssp. americanus. Within the haplotype M lineage, the introduced North American P. australis exhibited different biomass allocation patterns and resource/energy‐use strategies compared to its European ancestor group. A discriminant analysis of principal components separated the haplotype M and the haplotype E lineages completely along the first canonical axis, highly related to photosynthetic gas‐exchange parameters, photosynthetic energy‐use efficiency and payback time. The second canonical axis, highly related to photosynthetic nitrogen use efficiency and construction costs, significantly separated the introduced P. australis in North America from its European ancestor. Synthesis. We conclude that the European P. australis lineage was preadapted to be invasive prior to its introduction, and that the invasion in North America is further stimulated by rapid post‐introduction evolution in several advantageous traits. The multicomparison approach used in this study could be an effective approach for distinguishing preadaptation and post‐introduction evolution of invasive species. Further research is needed to link the observed changes in invasive traits to the genetic variation and the interaction with the environment.     

Modeling the role of the close‐range effect and environmental variables in the occurrence and spread of Phragmites australis in four sites on the Finnish coast of the Gulf of Finland and the Archipelago Sea

Phragmites australis, a native helophyte in coastal areas of the Baltic Sea, has significantly spread on the Finnish coast in the last decades raising ecological questions and social interest and concern due to the important role it plays in the ecosystem dynamics of shallow coastal areas. Despite its important implications on the planning and management of the area, predictive modeling of Phragmites distribution is not well studied. We examined the prevalence and progression of Phragmites in four sites along the Southern Finnish coast in multiple time frames in relation to a number of predictors. We also analyzed patterns of neighborhood effect on the expansion and disappearance of Phragmites in a cellular data model. We developed boosted regression trees models to predict Phragmites occurrences and produce maps of habitat suitability. Various Phragmites spread figures were observed in different areas and time periods, with a minimum annual expansion rate of 1% and a maximum of 8%. The water depth, shore openness, and proximity to river mouths were found influential in Phragmites distribution. The neighborhood configuration partially explained the dynamics of Phragmites colonies. The boosted regression trees method was successfully used to interpolate and extrapolate Phragmites distributions in the study sites highlighting its potential for assessing habitat suitability for Phragmites along the Finnish coast. Our findings are useful for a number of applications. With variables easily available, delineation of areas susceptible for Phragmites colonization allows early management plans to be made. Given the influence of reed beds on the littoral species and ecosystem, these results can be useful for the ecological studies of coastal areas. We provide estimates of habitat suitability and quantification of Phragmites expansion in a form suitable for dynamic modeling, which would be useful for predicting future Phragmites distribution under different scenarios of land cover change and Phragmites spatial configuration.     

The role of citrate on the oxidative metabolism of submerged cultures of Claviceps purpurea 129

In submerged cultures of Claviceps purpurea citrate utilization was stopped when the pool phosphate concentration decreased to 0.44–0.55 mmol per g of mycelial protein. The absence of citrate in the medium resulted in a 50% decrease of biomass, cessation of the alkaloid (agro-, elymo-, chanoclavine) synthesis, formation of a central large vacuole, cell enlargement and medium acidification by lactate. Citrate synthase and malate dehydrogenase gradually disappeared. Glucose-6-phosphate dehydrogenase was not detectable. The activities of glyceraldehyde-3-phosphate dehydrogenase and lactate dehydrogenase increased by one or two orders of magnitude higher than in control culture (with 80 mM citrate). Substrate competition on the pyruvate level is discussed.     

Effects of high Zn and Pb concentrations on Phragmites australis (Cav.) Trin. Ex. Steudel: Photosynthetic performance and metal accumulation capacity under controlled conditions

The response of Phragmites australis (Cav.) Trin. Ex. Steudel to zinc (Zn) and lead (Pb) was studied separately in two hydroponic tests, during a three weeks experiment. The effects on ecophysiology and biomass partitioning were evaluated during the metal treatments and at the recovery, and total metal content and accumulation capacity in different plant organs were assessed. Zn and Pb had different effects on the overall measured parameters, highlighting different mechanism of action. In particular, Zn concentration was higher in roots and, being a micronutrient, it was translocated into leaves, producing a reduction of assimilation rate, stomatal conductance (–71.9 and –81.3% respect to the control plant respectively), and a strong down regulation of photosystems functionality both at PSII and PSI level. Otherwise, Pb was accumulated mainly in the more lignified tissue such as rhizomes, with slightly effect on gas exchange. Chlorophyll a fluorescence highlighted that Pb inhibits the electron transfer process at the PSI donor side, without recovery after the removal of the metal stress. Despite these physiological limitations, P. australis showed a high capacity to accumulate both metals, and only slight reduction of biomass, being therefore a suitable species for phytoremediation interventions.     

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 (g_s), 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 (A_sat), maximum carboxylation rates of Rubisco (V_cmax), maximum electron transport rates (J_max), triose phosphate utilization rates (T_p), stomata conductance (g_s), as well as higher Rubisco carboxylation‐limited, RuBP regeneration‐limited and T_p‐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.     

氮沉降对黄河三角洲芦苇湿地土壤呼吸的影响

2012年6月至2012年10月, 对黄河三角洲芦苇(Phragmites australis)湿地进行了模拟氮沉降试验, 氮沉降水平分别为对照(CK, 0 kg N·hm^-2·a^-1)、低氮(LN, 50 kg N·hm^-2·a^-1)和高氮(HN, 100 kg N·hm^-2·a^-1)。利用LI-8100土壤碳通量测量系统测定土壤呼吸速率。结果表明, 氮沉降促进了芦苇湿地土壤呼吸作用, LN和HN处理使芦苇生长季(6-10月)平均土壤呼吸速率比CK分别提高19%和58%。积水改变了芦苇湿地土壤呼吸日动态。地面无积水时, 各处理土壤呼吸日动态均呈单峰型曲线; 地面有积水时, 土壤呼吸日动态峰值推后或无单峰型波动规律。积水影响土壤呼吸作用对温度的响应。地面无积水时, 各处理土壤呼吸速率均与气温呈极显著的正指数相关关系, 气温分别解释了CK、LN和HN处理下土壤呼吸季节变化的69.9%、64.5%和59.9%; 地面有积水时, 各处理土壤呼吸与气温相关性不显著。CK、LN和HN处理下土壤呼吸温度敏感性系数Q₁₀值分别为1.68、1.75和1.68, 表明LN处理增强了土壤呼吸温度敏感性, HN处理对其影响不显著。     

Invasion of Old World Phragmites australis in the New World: precipitation and temperature patterns combined with human influences redesign the invasive niche

After its introduction into North America, Euro‐Asian Phragmites australis became an aggressive invasive wetland grass along the Atlantic coast of North America. Its distribution range has since expanded to the middle, south and southwest of North America, where invasive P. australis has replaced millions of hectares of native plants in inland and tidal wetlands. Another P. australis invasion from the Mediterranean region is simultaneously occurring in the Gulf region of the United States and some countries in South America. Here, we analysed the occurrence records of the two Old World invasive lineages of P. australis (Haplotype M and Med) in both their native and introduced ranges using environmental niche models (ENMs) to assess (i) whether a niche shift accompanied the invasions in the New World; (ii) the role of biologically relevant climatic variables and human influence in the process of invasion; and (iii) the current potential distribution of these two lineages. We detected local niche shifts along the East Coast of North America and the Gulf Coast of the United States for Haplotype M and around the Mississippi Delta and Florida of the United States for Med. The new niche of the introduced Haplotype M accounts for temperature fluctuations and increased precipitation. The introduced Med lineage has enlarged its original subtropical niche to the tropics‐subtropics, invading regions with a high annual mean temperature (> ca. 10 °C) and high precipitation in the driest period. Human influence is an important factor for both niches. We suggest that an increase in precipitation in the 20th century, global warming and human‐made habitats have shaped the invasive niches of the two lineages in the New World. However, as the invasions are ongoing and human and natural disturbances occur concomitantly, the future distribution ranges of the two lineages may diverge from the potential distribution ranges detected in this study.     

Genetic diversity and gene flow estimates among three populations of Botryodiplodia theobromae causing die-back and bark canker of pear in Punjab

Three populations of Botryodiplodia theobromae causing die-back and bark canker of pear in Punjab were analysed to know about the genetic diversity, gene flow, heterogeneity and the probable rate of spread of races capable of overcoming the resistance present in elite cultivars. There was no relationship between the morphologically similar isolates and their pathogenic behaviour. Majority of the isolates of B. theobromae within and among the populations produced lesions of 1.9–7.2 × 0.8–3.1 cm size on detached pear twigs cv. Punjab Beauty. Incubation period and per cent infection also varied within and among the three populations of the pathogen. The genetic diversity was calculated on the basis of allele frequencies of nine simple sequence repeat (SSR) markers using Nei's genetic diversity formulae. Allele frequencies (X_ij ) varied significantly among the three populations ranging from 0.0 to 1.0. The mean genetic diversities within population (H _S) also varied in all the locations ranging from 0.18 to 0.36 indicating a high genetic diversity within the population. The total genetic diversity across all the populations (H _T) ranged between 0.1 and 0.5 with a mean of 0.36, and differed significantly from the mean H _S (0.25). Diversity indices of SSR loci varied from low (H _S < 0.1) to high (H _S > 0.4) across all the populations (G _ST) of B. theobromae in Punjab which were consistent over the presumed SSR loci with a mean G _ST of 0.29. The lower G _ST values of 0.13, 0.10, 0.11 and 0.13 were observed at LAS15-16, LAS27-28, BOT17-18 and BOT35-36 loci, respectively, showing high genetic diversity among all the isolates. However, at BOT19-20 locus, the G _ST value observed was 0.72 thereby indicating low diversity in the population at this locus. Out of 25 loci, 8 loci showed the gene flow (Nm) < 1 indicating a high genetic differentiation within the population and the remaining 6 loci showed the Nm > 1 indicating the frequent existence of gene flow across the populations of B. theobromae. Pairwise comparison (G _ST) values among all the loci were ranging from 0.18 to 0.19, which indicate a low genetic differentiation among the populations of B. theobromae. The high genetic diversity (H _S) within the population was observed in the present study suggesting that B. theobromae possess high variability in Punjab. Keeping this in view, new races would be expected soon outside the state of origin, as naturally occurring gene flow is likely to be increased by the human activity for comprehensive cultivation of pear in the near future. Therefore, it is difficult to predict the durability of resistant sources in Punjab, which in turn emphasises the identification and introgression of R genes into commercial cultivars.     

Effects of winter harvest on water and sediment chemistry in a stand of reed (Phragmites australis)

An investigation of the effects of repeated winter harvest and removal of dry reed culms on water and sediment chemistry was made in a shallow lake dominated by closed, monospecific reed stands. The oxygen concentration and redox potential in the water, as well as temperature, showed regular diurnal fluctuations inside the reed stands. Ammonium and nitrate + nitrite did not exhibit diurnal fluctuations and differences between a harvested and an unharvested reed area were small. Molybdate reactive phosphorus (MRP) concentration was higher in the unharvested area.Because the winter harvest caused an almost doubling of shoot production compared to the unharvested area, net addition of reed litter (leaves, stubble and culms) to the sediment of the harvested area was similar to that of the unharvested area. It is concluded that removal of reed culms in winter has the largest effect on microclimate in spring, and that the rate of mineralization and reducing intensity in the substrate are not immediately affected. Significantly higher summer standing crop of reed after harvest may have caused lower phosphate concentration in the water column and sediment interstitial water.