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1.  Preadaptation and post-introduction evolution facilitate the invasion of Phragmites australis in North America 
Ecology and Evolution  2014;4(24):4567-4577.
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.
doi:10.1002/ece3.1286
PMCID: PMC4278810  PMID: 25558352
Biomass allocation; common reed; common-environment experiment; discriminant analysis; ecophysiological trade-off; functional traits; invasion ecology; leaf construction cost; photosynthesis; standardized major axis (SMA)
2.  Expression of major photosynthetic and salt-resistance genes in invasive reed lineages grown under elevated CO2 and temperature 
Ecology and Evolution  2014;4(21):4161-4172.
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.
doi:10.1002/ece3.1282
PMCID: PMC4242567  PMID: 25505541
Common reed; Delta-type; EU-type; Mississippi River Delta; Phragmites australis; US Gulf Coast
3.  Response to multi-generational selection under elevated [CO2] in two temperature regimes suggests enhanced carbon assimilation and increased reproductive output in Brassica napus L. 
Ecology and Evolution  2013;3(5):1163-1172.
Functional plant traits are likely to adapt under the sustained pressure imposed by environmental changes through natural selection. Employing Brassica napus as a model, a multi-generational study was performed to investigate the potential trajectories of selection at elevated [CO2] in two different temperature regimes. To reveal phenotypic divergence at the manipulated [CO2] and temperature conditions, a full-factorial natural selection regime was established in a phytotron environment over the range of four generations. It is demonstrated that a directional response to selection at elevated [CO2] led to higher quantities of reproductive output over the range of investigated generations independent of the applied temperature regime. The increase in seed yield caused an increase in aboveground biomass. This suggests quantitative changes in the functions of carbon sequestration of plants subjected to increased levels of CO2 over the generational range investigated. The results of this study suggest that phenotypic divergence of plants selected under elevated atmospheric CO2 concentration may drive the future functions of plant productivity to be different from projections that do not incorporate selection responses of plants. This study accentuates the importance of phenotypic responses across multiple generations in relation to our understanding of biogeochemical dynamics of future ecosystems. Furthermore, the positive selection response of reproductive output under increased [CO2] may ameliorate depressions in plant reproductive fitness caused by higher temperatures in situations where both factors co-occur.
doi:10.1002/ece3.523
PMCID: PMC3678472  PMID: 23762504
Carbon dioxide; experimental evolution; global change; laboratory natural selection; oilseed rape; phenotypic divergence; plant-environment feedbacks
4.  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
5.  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
6.  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
7.  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
8.  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
9.  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
10.  Invasion strategies in clonal aquatic plants: are phenotypic differences caused by phenotypic plasticity or local adaptation? 
Annals of Botany  2010;106(5):813-822.
Background and Aims
The successful spread of invasive plants in new environments is often linked to multiple introductions and a diverse gene pool that facilitates local adaptation to variable environmental conditions. For clonal plants, however, phenotypic plasticity may be equally important. Here the primary adaptive strategy in three non-native, clonally reproducing macrophytes (Egeria densa, Elodea canadensis and Lagarosiphon major) in New Zealand freshwaters were examined and an attempt was made to link observed differences in plant morphology to local variation in habitat conditions.
Methods
Field populations with a large phenotypic variety were sampled in a range of lakes and streams with different chemical and physical properties. The phenotypic plasticity of the species before and after cultivation was studied in a common garden growth experiment, and the genetic diversity of these same populations was also quantified.
Key Results
For all three species, greater variation in plant characteristics was found before they were grown in standardized conditions. Moreover, field populations displayed remarkably little genetic variation and there was little interaction between habitat conditions and plant morphological characteristics.
Conclusions
The results indicate that at the current stage of spread into New Zealand, the primary adaptive strategy of these three invasive macrophytes is phenotypic plasticity. However, while limited, the possibility that genetic diversity between populations may facilitate ecotypic differentiation in the future cannot be excluded. These results thus indicate that invasive clonal aquatic plants adapt to new introduced areas by phenotypic plasticity. Inorganic carbon, nitrogen and phosphorous were important in controlling plant size of E. canadensis and L. major, but no other relationships between plant characteristics and habitat conditions were apparent. This implies that within-species differences in plant size can be explained by local nutrient conditions. All together this strongly suggests that invasive clonal aquatic plants adapt to a wide range of habitats in introduced areas by phenotypic plasticity rather than local adaptation.
doi:10.1093/aob/mcq176
PMCID: PMC2958791  PMID: 20826438
Alien weeds; biological invasion; clonal plants; Egeria densa; Elodea canadensis; establishment; genetic diversity; Lagarosiphon major; local adaptation; macrophytes; morphometric characters; phenotypic plasticity
11.  Nutrient and growth responses of cattail (Typha domingensis) to redox intensity and phosphate availability 
Annals of Botany  2009;105(1):175-184.
Background and Aims
In the Florida Everglades, the expansion of cattail (Typha domingensis) into areas once dominated by sawgrass (Cladium jamaicense) has been attributed to altered hydrology and phosphorus (P) enrichment. The objective of this study was to quantify the interactive effects of P availability and soil redox potential (Eh) on the growth and nutrient responses of Typha, which may help to explain its expansion.
Methods
The study examined the growth and nutrient responses of Typha to the interactive effects of P availability (10, 80 and 500 µg P L−1) and Eh level (−150, +150 and +600 mV). Plants were grown hydroponically in a factorial experiment using titanium (Ti3+) citrate as a redox buffer.
Key Results
Relative growth rate, elongation, root-supported tissue/root ratio, leaf length, lateral root length and biomass, as well as tissue nutrient concentrations, were all adversely affected by low Eh conditions. P availability compensated for the negative effect of low Eh for all these variables except that low P stimulated root length and nutrient use efficiency. The most growth-promoting treatment combination was 500 µg P L−1/ + 600 mV.
Conclusions
These results, plus previous data on Cladium responses to P/Eh combinations, document that high P availability and low Eh should benefit Typha more than Cladium as the growth and tissue nutrients of the former species responded more to excess P, even under highly reduced conditions. Therefore, the interactive effects of P enrichment and Eh appear to be linked to the expansion of Typha in the Everglades Water Conservation Area 2A, where both low Eh and enhanced phosphate availability have co-occurred during recent decades.
doi:10.1093/aob/mcp213
PMCID: PMC2794056  PMID: 19748907
Everglades; growth; nutrient; phosphorus; redox potential; Typha domingensis
12.  Genetic diversity in three invasive clonal aquatic species in New Zealand 
BMC Genetics  2010;11:52.
Background
Elodea canadensis, Egeria densa and Lagarosiphon major are dioecious clonal species which are invasive in New Zealand and other regions. Unlike many other invasive species, the genetic variation in New Zealand is very limited. Clonal reproduction is often considered an evolutionary dead end, even though a certain amount of genetic divergence may arise due to somatic mutations. The successful growth and establishment of invasive clonal species may be explained not by adaptability but by pre-existing ecological traits that prove advantageous in the new environment. We studied the genetic diversity and population structure in the North Island of New Zealand using AFLPs and related the findings to the number of introductions and the evolution that has occurred in the introduced area.
Results
Low levels of genetic diversity were found in all three species and appeared to be due to highly homogeneous founding gene pools. Elodea canadensis was introduced in 1868, and its populations showed more genetic structure than those of the more recently introduced of E. densa (1946) and L. major (1950). Elodea canadensis and L. major, however, had similar phylogeographic patterns, in spite of the difference in time since introduction.
Conclusions
The presence of a certain level of geographically correlated genetic structure in the absence of sexual reproduction, and in spite of random human dispersal of vegetative propagules, can be reasonably attributed to post-dispersal somatic mutations. Direct evidence of such evolutionary events is, however, still insufficient.
doi:10.1186/1471-2156-11-52
PMCID: PMC2902404  PMID: 20565861
13.  Can differences in phosphorus uptake kinetics explain the distribution of cattail and sawgrass in the Florida Everglades? 
BMC Plant Biology  2010;10:23.
Background
Cattail (Typha domingensis) has been spreading in phosphorus (P) enriched areas of the oligotrophic Florida Everglades at the expense of sawgrass (Cladium mariscus spp. jamaicense). Abundant evidence in the literature explains how the opportunistic features of Typha might lead to a complete dominance in P-enriched areas. Less clear is how Typha can grow and acquire P at extremely low P levels, which prevail in the unimpacted areas of the Everglades.
Results
Apparent P uptake kinetics were measured for intact plants of Cladium and Typha acclimated to low and high P at two levels of oxygen in hydroponic culture. The saturated rate of P uptake was higher in Typha than in Cladium and higher in low-P acclimated plants than in high-P acclimated plants. The affinity for P uptake was two-fold higher in Typha than in Cladium, and two- to three-fold higher for low-P acclimated plants compared to high-P acclimated plants. As Cladium had a greater proportion of its biomass allocated to roots, the overall uptake capacity of the two species at high P did not differ. At low P availability, Typha increased biomass allocation to roots more than Cladium. Both species also adjusted their P uptake kinetics, but Typha more so than Cladium. The adjustment of the P uptake system and increased biomass allocation to roots resulted in a five-fold higher uptake per plant for Cladium and a ten-fold higher uptake for Typha.
Conclusions
Both Cladium and Typha adjust P uptake kinetics in relation to plant demand when P availability is high. When P concentrations are low, however, Typha adjusts P uptake kinetics and also increases allocation to roots more so than Cladium, thereby improving both efficiency and capacity of P uptake. Cladium has less need to adjust P uptake kinetics because it is already efficient at acquiring P from peat soils (e.g., through secretion of phosphatases, symbiosis with arbuscular mycorrhizal fungi, nutrient conservation growth traits). Thus, although Cladium and Typha have qualitatively similar strategies to improve P-uptake efficiency and capacity under low P-conditions, Typha shows a quantitatively greater response, possibly due to a lesser expression of these mechanisms than Cladium. This difference between the two species helps to explain why an opportunistic species such as Typha is able to grow side by side with Cladium in the P-deficient Everglades.
doi:10.1186/1471-2229-10-23
PMCID: PMC2923519  PMID: 20141632

Results 1-13 (13)