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1.  Interactive effects of elevated temperature and CO2 on two phylogeographically distinct clones of common reed (Phragmites australis) 
AoB Plants  2013;5:pls051.
One European and one Mediterranean Phragmites australis genotype (DK clone and ALG clone, respectively) showed distinct aboveground growth and physiology in response to different treatment combinations of elevated CO2 and temperature according to their genetic background. The DK clone was the most responsive clone.
The aboveground growth, physiological and biochemical parameters of two clones of the cosmopolitan wetland grass Phragmites australis, grown at four treatment combinations of temperature and CO2, were investigated to elucidate whether their climate response differed due to inherent differences in their ecological adaptation. The two phylogeographically distinct P. australis clones (DK clone, European genetic background; ALG clone, Mediterranean genetic background) were grown for 151 days in phytotrons at 19/12 °C (day/night temperature) and 390 ppm CO2, and at elevated temperature (+5 °C) and CO2 (700 ppm) with treatment factors alone or in combination. The ALG clone had 2–4 times higher aboveground biomass, higher light-saturated rates of photosynthesis (Pmax), maximum electron transport rates (ETRmax) and Rubisco activity, and higher photosynthetic nitrogen-use efficiency than the DK clone. The DK clone, however, produced more shoots, leaves and side-shoots, and had 9–51 % higher specific leaf area and 15–39 % higher leaf nitrogen concentration than the ALG clone. Although elevated atmospheric CO2 alone barely affected the aboveground growth of the two P. australis clones, simultaneously elevated temperature and CO2 stimulated growth and aboveground biomass. Overall, elevated CO2 stimulated photosynthesis, but the clones responded differently to a concomitant increase in CO2 and temperature, depending on the phylogeographic background of the plant. The DK clone showed overall stronger responses, and can be considered the more plastic of the two clones with respect to CO2 and temperature. Thus, the DK clone may be better adapted to climate change than the ALG clone, at least in the short term.
doi:10.1093/aobpla/pls051
PMCID: PMC4104621
Algeria; climate change; Denmark; Mediterranean Phragmites; RERAF phytotron; temperate Phragmites
2.  Differences in salinity tolerance of genetically distinct Phragmites australis clones 
AoB Plants  2013;5:plt019.
The common reed (Phragmites australis) is a clonal wetland grass with high genetic variability. Clone-specific differences are reflected in morphological and physiological traits, and hence in the ability to cope with environmental stress. The responses to progressively increasing salinity of fifteen distinct Phragmites australis clones reveal genotype-related strategies of salt avoidance and exclusion. The salinity-induced inhibition in shoot elongation rate and photosynthesis varies widely between clones. The differences can be partially attributed to their geographic range, but not correlated to ploidy level. Thus, the genetic background is a major factor influencing the salinity tolerance of distinct Phragmites australis clones.
Different clones of the wetland grass Phragmites australis differ in their morphology and physiology, and hence in their ability to cope with environmental stress. We analysed the responses of 15 P. australis clones with distinct ploidy levels (PLs) (4n, 6n, 8n, 10n, 12n) and geographic origins (Romania, Russia, Japan, Czech Republic, Australia) to step-wise increased salinity (8, 16, 24, 32, 40, 56 and 72 ppt). Shoot elongation rate, photosynthesis and plant part-specific ion accumulation were studied in order to assess if traits associated with salinity tolerance can be related to the genetic background and the geographic origin of the clones. Salt stress affected all clones, but at different rates. The maximum height was reduced from 1860 mm in control plants to 660 mm at 40 ppt salinity. The shoot elongation rate of salt-exposed plants varied significantly between clones until 40 ppt salinity. The light-saturated photosynthesis rate (Pmax) was stimulated by a salinity of 8 ppt, but decreased significantly at higher salinities. The stomatal conductance (gs) and the transpiration rate (E) decreased with increasing salinity. Only three clones survived at 72 ppt salinity, although their rates of photosynthesis were strongly inhibited. The roots and basal leaves of the salt-exposed plants accumulated high concentrations of water-extractable Na+ (1646 and 1004 µmol g−1 dry mass (DM), respectively) and Cl− (1876 and 1400 µmol g−1 DM, respectively). The concentrations of water-extractable Mg2+ and Ca2+ were reduced in salt-exposed plants compared with controls. The variation of all the measured parameters was higher among clones than among PLs. We conclude that the salinity tolerance of distinct P. australis clones varies widely and can be partially attributed to their longitudinal geographic origin, but not to PL. Further investigation will help in improving the understanding of this species' salt tolerance mechanisms and their connection to genetic factors.
doi:10.1093/aobpla/plt019
PMCID: PMC4104622
Common reed; ecophysiology; geographic range; ion concentration; ploidy level; salt-stress tolerance
3.  Photosynthesis of co-existing Phragmites haplotypes in their non-native range: are characteristics determined by adaptations derived from their native origin? 
AoB Plants  2013;5:plt016.
Several Phragmites lineages differing in origin and phenotype co-exist in the Gulf Coast of North America. We collected rhizomes of four lineages and propagated them in a common environment to compare photosynthetic characteristics. We observed substantial differences among and within lineages. As the lineages originating in Africa and in the Mediterranean region had higher photosynthetic capacity than the lineages originating in Eurasia, and showed typical ecophysiological traits of plants adapted to warm and arid climates, we concluded that the differences observed are due to adaptations acquired in the native ranges. The four lineages can therefore be regarded as ecotypes.
The Gulf Coast of North America (GC) is a ‘hot spot’ of Phragmites diversity as several lineages (defined according to the haplotypes of their chloroplast DNA) differing in origin, genetic traits and phenotype co-exist and interbreed in this area. We analysed differences in photosynthetic characteristics among and within four haplotypes to understand if differences in gas exchange can be attributed to adaptations acquired in their native ranges. We collected rhizomes of four GC haplotypes (I2, M1, M and AI; including the phenotypes ‘Land-type’, ‘Delta-type’, ‘EU-type’ and ‘Greeny-type’) and propagated them in a common controlled environment to compare photosynthesis–irradiance responses, CO2 responses, chlorophyll fluorescence, the activity of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), specific leaf area (SLA), pigment contents, stomatal density and guard cell length. The maximum light-saturated photosynthetic rate, Amax, in the haplotype I2 (Land-type) and haplotype M1 (Delta-type) (34.3–36.1 µmol CO2 m−2 s−1) was higher than that in the invasive Eurasian haplotype M (22.4 ± 2.3 µmol CO2 m−2 s−1). The Amax of haplotype AI (Greeny3-type) was 29.1 ± 4.0 µmol CO2 m−2 s−1 and did not differ from the Amax of the other haplotypes. The carboxylation rate (Vcmax) and electron transport rate (Jmax) followed the same pattern as Amax. The haplotypes also differed in SLA (17.0–24.3 m2 kg−1 dry mass) and pigment content, whereas stomatal density and guard cell length, Rubisco activity and chlorophyll fluorescence did not differ significantly among haplotypes. The high photosynthetic activity and gas-exchange capacity of the two haplotypes originating in tropical Africa and the Mediterranean area (haplotypes I2 and M1) are apparently adaptations derived from their native ranges. Hence, the haplotypes can be regarded as ecotypes. However, it remains unclear how these differences relate to plant competitiveness and fitness in the GC of North America environment.
doi:10.1093/aobpla/plt016
PMCID: PMC4104645
Adaptations; Gulf Coast of North America; genotypes; haplotypes; invasion; photosynthesis; Phragmites
4.  Phenotypic traits of Phragmites australis clones are not related to ploidy level and distribution range 
AoB Plants  2012;2012:pls017.
The present study reveals significant genetically determined differences in a range of growth and ecophysiological traits between different Phragmites australis genotypes, and provides evidence that the differences are neither related to ploidy level per se nor to the phylogeographic relationships of the genotypes.
Background and aims
Phragmites australis is a wetland grass with high genetic variability, augmented by its cosmopolitan distribution, clonal growth form and large variation in chromosome numbers. Different ploidy levels and ecotypes differ in morphology and ecophysiological traits, and may possess different levels of phenotypic variation. The aim of this study was to quantify the natural variation in ecophysiological characteristics of P. australis, and to explore whether differences in ecophysiological traits can be related to ploidy levels or to the geographic origin of the clones.
Methodology
Fifteen clones of P. australis from Europe and Asia/Australia, representing five ploidy levels (4x, 6x, 8x, 10x and 12x), were grown in a common garden design for 119 days. Plant growth and light-saturated rate of photosynthesis (Pmax), stomatal conductance (gs), water use efficiency (WUE) and concentrations of photosynthetic pigments and mineral ions in the leaves were measured.
Principal results
The growth of the plants and most ecophysiological parameters differed significantly between clones. The mean maximum shoot height varied from 0.9 to 1.86 m, Pmax from 9.7 to 27 µmol m−2 s−1, gs from 0.22 to 1.41 mol m−2 s−1 and WUE from 13 to 47 µmol mol−1. The concentrations of chlorophylls did not vary significantly between clones, but the chlorophyll a/b ratio and the concentrations of total carotenoids did. The observed differences were not explained either by the ploidy level per se or by the geographic origin or phylogenetic relationships of the clones.
Conclusions
Phylogeographic relationships in P. australis on a global scale do not mirror the environment where the adaptations have evolved, and high phenotypic variation among and within clones complicates comparative studies. Future studies aimed at explaining differences in plant behaviour between P. australis populations should be careful in the selection of target genotypes and/or populations, and should avoid generalizing their findings beyond the genotypes and/or populations studied.
doi:10.1093/aobpla/pls017
PMCID: PMC3407373  PMID: 22848787
5.  Exploring the borders of European Phragmites within a cosmopolitan genus 
AoB Plants  2012;2012:pls020.
European Phragmites australis is one of four main cp-DNA haplotype clusters present worldwide. The European gene pool extends from North America to Far East Asia and South Africa. Extensive gene flow occurs only within the temperate region of Europe.
Background and aims
Two Phragmites australis taxa are recognized in Europe: P. australis ssp. altissimus, also known as Phragmites isiaca, in the Mediterranean region and P. australis in the temperate region. Another taxonomic group in the Mediterranean is Phragmites frutescens. European genotypes are diverse genetically, cytologically and morphologically, and are related to African, Asiatic and American genotypes. We investigated chloroplast DNA (cpDNA) diversity in Europe and defined the current borders of the European gene pool.
Methodology
We analysed chloroplast variation with parsimony and genetic distance methods, and compared it with that of nuclear amplified fragment length polymorphism and microsatellites. We also investigated the phenological pattern of 188 genotypes collected worldwide in a common garden in Denmark. We assumed that non-flowering genotypes could indicate climatic, geographic and/or reproductive barriers to dispersal and would have been recorded in the genetic pattern as groups genetically isolated from, or within, the European pool.
Principal results
The European P. australis gene pool extends from North America to the Far East and South Africa. However, African and North American genotypes are differentiating from the European genotypes. Mediterranean P. australis is genetically different from temperate P. australis and shares several similarities with Phragmites mauritianus in Africa and Phragmites karka in Asia. Phragmites frutescens shares the cpDNA sequences with both these tropical species. Two DNA bands can distinguish Mediterranean P. australis from P. frutescens and P. mauritianus and from temperate P. australis, and reveal possible hybrids among these species in the Mediterranean region. Phenological data confirmed possible gene flow within the temperate region of Europe, whereas the Mediterranean genotypes did not set inflorescences in Denmark, suggesting reproductive barriers between temperate and Mediterranean P. australis.
Conclusions
European P. australis appears as one of four main Phragmites groups known in the world. Further research is needed to understand the implications of long-distance dispersal at the population level.
doi:10.1093/aobpla/pls020
PMCID: PMC3435523  PMID: 22962631
6.  Regression analysis of growth responses to water depth in three wetland plant species 
AoB Plants  2012;2012:pls043.
Variability in plant flooding tolerance is often associated with differential growth responses to increasing water depth. This study highlights how morphological responses conferring flooding tolerance differ, using non-linear and quantile regression to quantitatively compare flooding-related growth responses of three species.
Background and aims
Plant species composition in wetlands and on lakeshores often shows dramatic zonation, which is frequently ascribed to differences in flooding tolerance. This study compared the growth responses to water depth of three species (Phormium tenax, Carex secta and Typha orientalis) differing in depth preferences in wetlands, using non-linear and quantile regression analyses to establish how flooding tolerance can explain field zonation.
Methodology
Plants were established for 8 months in outdoor cultures in waterlogged soil without standing water, and then randomly allocated to water depths from 0 to 0.5 m. Morphological and growth responses to depth were followed for 54 days before harvest, and then analysed by repeated-measures analysis of covariance, and non-linear and quantile regression analysis (QRA), to compare flooding tolerances.
Principal results
Growth responses to depth differed between the three species, and were non-linear. Phormium tenax growth decreased rapidly in standing water >0.25 m depth, C. secta growth increased initially with depth but then decreased at depths >0.30 m, accompanied by increased shoot height and decreased shoot density, and T. orientalis was unaffected by the 0- to 0.50-m depth range. In P. tenax the decrease in growth was associated with a decrease in the number of leaves produced per ramet and in C. secta the effect of water depth was greatest for the tallest shoots. Allocation patterns were unaffected by depth.
Conclusions
The responses are consistent with the principle that zonation in the field is primarily structured by competition in shallow water and by physiological flooding tolerance in deep water. Regression analyses, especially QRA, proved to be powerful tools in distinguishing genuine phenotypic responses to water depth from non-phenotypic variation due to size and developmental differences.
doi:10.1093/aobpla/pls043
PMCID: PMC3526336  PMID: 23259044

Results 1-6 (6)