Effects of arbuscular mycorrhizae on biomass and nutrients in the aquatic plant Littorella uniflora

1. It has been hypothesised that the symbiosis with arbuscular mycorrhizal fungi (AMF) leads to a higher uptake of phosphorus (P) and nitrogen (N) in aquatic plants, but it has never been shown experimentally without the use of fungicides. In particular, the symbiosis may be important for nutrient uptake by isoetids in oligotrophic lakes, where low concentrations of inorganic N and P both in the water and in the sediment limit the growth of plants and where symbiosis facilitates the uptake of nutrients from the sediment. 2. Plants of the isoetid Littorella uniflora were propagated under the sterile conditions without an AMF infection. The plants were then grown for 60 days with and without re‐infection by AMF, and with either high (150 μm ) or low (ambient concentration approximately 15 μm ) CO₂ concentration. 3. The study proved that the symbiosis between AMF and L. uniflora had a positive impact on the retention of N and P in the plants at very low nutrient concentrations in the water and on biomass development. Shoot biomass and standing stocks of both P and N were significantly higher in re‐infected plants. 4. Raised CO₂ concentration resulted in a fivefold increase in hyphal infection, but had no impact on the number of arbuscules and vesicles in the cross sections. There were significantly higher biomass and lower tissue P and N concentrations in the plants from high CO₂ treatments. This resulted in similar standing stocks of P and N in plants from low and high CO₂ treatments. 5. The results from this study showed that the symbiosis between AMF and L. uniflora is an important adaptation enabling isoetids to grow on nutrient‐poor sediments in oligotrophic lakes.     

Effects of CO2 concentration on growth of filamentous algae and Littorella uniflora in a Danish softwater lake

An experiment was conducted from May to November in Lake Hampen, Denmark, to study the effect of higher CO₂ concentration on the biomass of filamentous algae. Three enclosures (1.5 m diameter) were enriched with free CO₂ to ∼10 times atmospheric equilibrium (∼170 μM) and three enclosures were kept at atmospheric equilibrium (∼17 μM). The isoetid Littorella uniflora dominated the vegetation in the enclosures. Low concentrations of nitrate and phosphate in the water were observed, especially in the summer months. During the summer, a high biomass of filamentous algae (dominated by Zygnema sp.) developed in both types of enclosures (18–58 g dry wt. m⁻² in July and August), but the biomass of algae was significantly higher (1.9–38 times) in the CO₂ enriched enclosures than in enclosures with low CO₂ concentration. L. uniflora biomass, especially leaf biomass, also showed a significant positive response to increased CO₂ concentration (75.0 ± 10.4 and 133.3 ± 42.5 g dry wt. m⁻² at low and high CO₂ concentrations, respectively) even though the massive filamentous algal growth decreased the light intensity. Both filamentous algae (in August) and L. uniflora showed lower tissue concentrations of N and P at high CO₂ concentration.     

Competition between isoetids and invading elodeids at different concentrations of aquatic carbon dioxide

1. The growth of submerged macrophytes in softwater lakes is often assumed to be carbon limited. Isoetid species are well adapted to grow at low carbon availability and therefore commonly dominate the submerged macrophyte vegetation in softwater lakes. In many such lakes, however, large‐scale invasions of fast‐growing elodeid species, replacing the isoetid vegetation, have been observed. 2. In a laboratory experiment, we tested how rising aquatic carbon availability, in interaction with different densities of the isoetid Littorella uniflora, affected the growth (and thereby the potential invasion success) of the elodeid Myriophyllum alterniflorum. For this purpose, the growth of M. alterniflorum was determined at a combination of three concentrations of dissolved CO₂ (15, 90, 200 μmol L^?1) and three densities of L. uniflora (0, 553, 1775 plants m^?2). 3. At an ambient CO₂ of 15 μmol L^?1, M. alterniflorum could not sustain itself, whereas at raised CO₂ concentrations, growth became positive and increased with higher CO₂ availability. 4. The presence of L. uniflora, independent of its density, reduced the growth of M. alterniflorum by 50%. Whether this is related to nutrient availability or other factors is not clear. 5. Despite the growth reduction of M. alterniflorum by L. uniflora, at CO₂ ≥90 μmol L^?1, L. uniflora was still overgrown by M. alterniflorum. This may imply that, in field situations, M. alterniflorum can invade softwater systems with relatively high CO₂ availability, even in the presence of dense stands of L. uniflora.     

Long‐term effects of liming in Norwegian softwater lakes: the rise and fall of bulbous rush (Juncus bulbosus) and decline of isoetid vegetation

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Groundwater seepage stimulates the growth of aquatic macrophytes

1. The impact of groundwater seepage on the growth of submerged macrophytes was investigated in experiments on the isoetid Littorella uniflora and the elodeid Myriophyllum alterniflorum both in the laboratory and in the field. Isoetids rely mostly on sediment‐derived CO₂ and nutrients via root uptake, whereas elodeids acquire their inorganic carbon and nutrients from the water column. We thus hypothesised that L. uniflora would respond positively to seeping ground water as it should improve both CO₂ and nutrient supply. 2. Laboratory experiments were conducted by percolating vegetated cores containing natural sediment or technical sand with artificial ground water of high CO₂ concentrations and with either high or low levels of nutrients. Field experiments were conducted in the oligotrophic Lake Hampen, Denmark, with custom‐built seepage‐growth chambers that permitted a near‐natural flow‐through of seeping ground water. Chambers with a solid bottom, and thus no flow‐through of seeping ground water, served as controls in both laboratory and field experiments. In the field, seepage chambers were installed at a site with relatively high seepage fluxes (ground water from forest catchment), at a site with much lower seepage fluxes but with higher nutrient concentrations (ground water from agricultural catchment) and at a reference site with no net discharge or recharge of ground water. 3. Positive growth responses were observed in the field at transects with high groundwater discharge compared to the control chambers with no seepage. No growth response was observed at the reference transect with low or alternating direction of groundwater seepage. The growth rates of L. uniflora in the field were significantly higher in seepage treatments compared to control treatments, and final plant mass was up to 70% higher than that for plants where seepage was excluded. In areas with high groundwater discharge, a strong positive correlation was found between groundwater seepage fluxes, growth rates, and final plant mass for L. uniflora, while there was no such relationship at the reference transect. The growth of M. alterniflorum was also significantly affected by groundwater seepage, but to a lesser degree than L. uniflora. Laboratory experiments generally showed the same trend for both L. uniflora and M. alterniflorum, and the positive influence of seeping ground water was apparently related to increased inorganic carbon supply and, to a lesser degree, improved nutrient availability. 4. Groundwater discharge results in enhanced growth of isoetids and to some extent elodeids inhabiting a groundwater‐fed softwater lake. We propose that the shallow dense vegetation present where most of the discharge takes place acts as a biological filter that retains nutrients that otherwise would end up in the water column and could result in increased algal growth.     

CAM-Like carbon pathway in submerged aquatic plants

CAM-like photosynthesis was found in the isoetid aquatic plantsLittorella uniflora andIsoetes lacustris, but not in the isoetid speciesLobelia dortmanna or in the elodeidElodea canadensis. Of the taxa studied, the first three are known to utilize sediment-borne CO₂, whereasElodea is dependent on bicarbonate.     

Epiphytic algae and macroinvertebrates on submerged and floating-leaved macrophytes in an Italian lake

1. The presence of contiguous beds of submerged (Myriophyllum spicatum, Ceratophyllum demersum and Najas marina) and floating-leaved (Trapa natans) vegetation in a north Italian lake allowed us to test the effect of the different host architecture on epiphytic algae and invertebrates and to predict the consequences for the lake of changes in the predominant vegetation. 2. Epiphyton development, measured as carbon, nitrogen, phosphorus, chlorophyll a (Chl a), phaeophytin and as algal and macroinvertebrate density, was significantly higher on submerged plants than on T. natans. The C : Chl a ratio, a proxy of the ratio of heterotrophs to autotrophs, was higher on the floating-leaved plants. The elemental (C : N : P) and pigment (Chl a : phaeophytin) ratios were not significantly different between the two vegetation types. 3. The taxonomic composition of epiphytic algae and invertebrates was similar on the different plants. The more varied morphology of the floating-leaved T. natans resulted in a higher diversity of epiphytic algae, however, but not of macroinvertebrates. 4. There was a significant inverse relationship between epiphyton biomass and the standing crop of the host plant, suggesting a key role for light and water exchange in epiphyton development. 5. Replacement of floating-leaved by submerged plants would increase the total biomass of epiphytic algae and invertebrates.     

Influence of quantity and lability of sediment organic matter on the biomass of two isoetids,Littorella uniflora and Echinodorus repens

1. Despite real improvement in the water quality of many previously eutrophic lakes, the recovery of submerged vegetation has been poor. This lack of recovery is possibly caused by the accumulation of organic matter on the top layer of the sediment, which is produced under eutrophic conditions. Hence, our objective was to study the combined effects of quantity and lability of sediment organic matter on the biomass of Echinodorus repens and Littorella uniflora and on the force required to uproot plants of L. uniflora. 2. Lake sediments, rich in organic matter, were collected from four lakes, two with healthy populations of isoetids and two from which isoetids had disappeared. The four lake sediments were mixed with sand to prepare a range of experimental sediments that differed in quantity and lability of sediment organic matter. Two isoetid species, E. repens and L. uniflora, were grown in these sediments for 8 weeks. Sediment quality parameters, including elemental composition, nutrient availability and mineralisation rates, were determined on the raw sources of sediment from the lakes. Porewater and surface water were analysed for the chemical composition in all mixtures. At the end of the experiment, plants were harvested and their biomass, tissue nutrient concentration and (for L. uniflora) uprooting force were measured. 3. For both species, all plants survived and showed no signs of stress on all types of sediment. The biomass of E. repens increased as the fraction of organic matter was increased (from 6 to 39% of organic content, depending upon sediment type). However, in some of the sediment types, a higher fraction of organic matter led to a decline in biomass. The biomass of L. uniflora was less responsive to organic content and was decreased significantly only when the least labile sediment source was used to create the gradient of organic matter. The increase in shoot biomass for both species was closely related to higher CO₂ concentrations in the porewater of the sediment. The force required to uproot L. uniflora plants over a range of sediment organic matter fitted a Gaussian model; it reached a maximum at around 15% organic matter and declined significantly above that. 4. Increasing organic matter content of the sediment increased the biomass of isoetid plants, as the positive effects of higher CO₂ production outweighed the negative effects of low oxygen concentration in more (labile) organic sediments. However, sediment organic matter can adversely affect isoetid survival by promoting the uprooting of plants.     

Physiological and photosynthetic plasticity in the amphibious, freshwater plant, Littorella uniflora, during the transition from aquatic to dry terrestrial environments

The physiological and photosynthetic responses of Littorella uniflora (L.) Ascherson, an amphibious macrophyte of isoetid life form, to rapid and prolonged emersion onto dry land, was studied at a reservoir. Water relations were little affected in the short term, but declining water potential and turgor pressure indicated water stress after flowering. High leaf lacunal CO₂ concentrations suggested continued CO₂ uptake from sediments. In contrast, a switch from Crassulacean acid metabolism (CAM) to C₃ photosynthesis was indicated by much lower levels of ΔH⁺ (down minus dusk titratable acidity) and phosphoenolpyruvate carboxylase (PEPC) activity in new terrestrial leaves, 7–8‐fold higher activity of ribulose bisphosphate carboxylase oxygenase (Rubisco), and increased chlorophyll and soluble protein contents. Accumulated nitrate and amino acid pools were depleted, whereas storage of carbohydrates as soluble sugars, fructan and starch increased. Plant carbon and nitrogen isotope ratios (δ¹³C and δ¹⁵N) declined, perhaps reflecting changes in C fixation processes, N metabolism, and source C and N. In leaves of plants grown half‐emersed for an extended period, contrasting activities of PEPC and Rubisco were found in submersed and emersed portions. Overall, L. uniflora showed considerable phenotypic plasticity, yet seemed to remain poised for re‐submersion; these characteristics could be adaptive in the unpredictable water margin habitat.     

Water‐level fluctuations affect sediment properties,carbon flux and growth of the isoetid Littorella uniflora in oligotrophic lakes

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Effects of elevated CO2 and vascular plants on evapotranspiration in bog vegetation

We determined evapotranspiration in three experiments designed to study the effects of elevated CO₂ and increased N deposition on ombrotrophic bog vegetation. Two experiments used peat monoliths with intact bog vegetation in containers, with one experiment outdoors and the other in a greenhouse. A third experiment involved monocultures and mixtures of Sphagnum magellanicum and Eriophorum angustifolium in containers in the same greenhouse. To determine water use of the bog vegetation in July–August for each experiment and each year we measured water inputs and outputs from the containers. We studied the effects of elevated CO₂ and N supply on evapotranspiration in relation to vascular plant biomass and exposure of the moss surface (measured as height of the moss surface relative to the container edge). Elevated CO₂ reduced water use of the bog vegetation in all three experiments, but the CO₂ effect on evapotranspiration interacted with vascular plant biomass and exposure of the moss surface. Evapotranspiration in the outdoor experiment was largely determined by evaporation from the Sphagnum moss surface (as affected by exposure to wind) and less so by vascular plant transpiration. Nevertheless, elevated CO₂ significantly reduced evapotranspiration by 9–10% in the outdoor experiment. Vascular plants reduced evapotranspiration in the outdoor experiment, but increased water use in the greenhouse experiments. The relation between vascular plant abundance and evapotranspiration appears to depend on wind conditions; suggesting that vascular plants reduce water losses mainly by reducing wind speed at the moss surface. Sphagnum growth is very sensitive to changes in water level; low water availability can have deleterious effects. As a consequence, reduced evapotranspiration in summer, whether caused by elevated CO₂ or by small increases in vascular plant cover, is expected to favour Sphagnum growth in ombrotrophic bog vegetation.     

Oxygen stress and reduced growth of Lobelia dortmanna in sandy lake sediments subject to organic enrichment

1. Lobelia dortmanna is a common representative of the small isoetid plants dominating the vegetation in nutrient‐poor lakes in Europe and North America. Because of large permeable root surfaces and continuous air lacunae Lobelia exchanges the majority of O₂ and CO₂ during photosynthesis across the roots. This leads to profound diel pulses of O₂ and CO₂ in sandy sediments with low microbial O₂ consumption rates. The ready radial root loss of O₂ may, however, make Lobelia very susceptible to more reducing sediments. Therefore, we grew Lobelia for 6 months on natural and organically enriched sandy sediments to test how: (i) root oxygenation influenced degradation of organic matter and depth profiles of N and C; (ii) Lobelia and microbial O₂ consumption rates influenced pool size and depth penetration of O₂ in the sediments; and (iii) sediment enrichment influenced growth and mineral nutrition of Lobelia. 2. Naturally low‐organic sediments (0.32% DW) accumulated organic C and N during the experiment as a result of growth of Lobelia and surface micro‐algae. In contrast, surface layers of enriched sediments (0.58, 0.87 and 2.46% DW) lost organic C and N because of enhanced mineralisation rates because of oxygen availability. In deeper layers of enriched sediments no significant differences in organic C and N pools were found between plant‐covered and plant‐free sediments probably because faster organic degradation because of root oxygenation was balanced by release of organic matter from the plants and because short roots with dense Fe‐Mn coatings in the most enriched sediments constrained O₂ release. 3. Depth‐integrated O₂ pools were much higher in light than darkness, higher in plant‐covered than plant‐free sediments and higher in sandy than in organically enriched sediments. All sediments had a primary O₂ maximum 1–2 mm below the sediment surface in light because of photosynthesis of micro‐algae. Plant‐covered sediments of low organic content (0.32 and 0.58% DW) also had a secondary deep maximum (2–4 cm) because of higher O₂ release from Lobelia roots than microbial O₂ consumption. Nitrification occurred here resulting in depletion of NH and accumulation of NO. In low organic sediments, oxygen pools increased with higher plant biomass both in light and darkness. The deep O₂ and NO₃ maxima disappeared in high organic sediments of greater O₂ consumption rates and smaller O₂ release rates. 4. Lobelia was stressed by increasing O₂ consumption rate of the sediments. Plant weight and leaf number declined twofold and maximum root length declined fourfold suggesting severe problems maintaining sufficient axial O₂ transport to the root tips because of rapid radial O₂ loss. Despite markedly higher nutrient concentrations in the enriched sediments, leaf‐N declined twofold and leaf‐P declined fourfold to growth‐limiting levels. These responses can be explained by constrains on mycorrhisal activity, root metabolism and vascular transport because of O₂ depletion. Management efforts to stop the decline and ensure the recovery of the isoetid vegetation should therefore focus on improving water quality as well as sediment suitability for growth.     

Could rising aquatic carbon dioxide concentrations favour the invasion of elodeids in isoetid‐dominated softwater lakes?

1. During the past century, isoetid vegetation types in softwater lakes have often been invaded by faster‐growing elodeids. In these C‐limited systems, this may be related to rising aquatic CO₂ levels. 2. In a laboratory experiment we tested the growth response of two elodeid species, Myriophyllum alterniflorum and Callitriche hamulata, at four different CO₂ levels, ranging from 20 to 230 μmol L⁻¹. In addition, we tested the effect of the nutrient status of the sediment on the growth of C. hamulata at the different CO₂ levels. 3. Shoot and root growth increased with rising CO₂ availability. Irrespective of sediment type, growth was minimal to negative at the lowest CO₂ treatment level, while becoming positive at CO₂ levels around 40–50 μmol L⁻¹. Substantial growth was only obtained when the macrophytes were growing on mesotrophic sediments. The plants reached close to maximal growth at CO₂ levels of c. 100 μmol L⁻¹. 4. Within this experiment, the growth of C. hamulata at CO₂ levels above 90 μmol L⁻¹ may have been limited by N and P availability in both sediment types. The growth rate of M. alterniflorum did not seem to be limited by N and P availability, most likely due to its much higher relative root production. 5. The experimental results show that neither M. alterniflorum nor C. hamulata is able to invade isoetid‐dominated softwater lakes at very low aquatic CO₂ concentrations. However, if the sediments contain enough nutrients, a rise in aquatic CO₂ could allow the invasion of elodeid species leading to the subsequent disappearance of slow‐growing isoetids.     

A multidisciplinary approach to understanding the recent and historical occurrence of the freshwater plant, Littorella uniflora

1. The vulnerability of softwater, oligotrophic lakes to eutrophication has caused the disappearance of many, if not most, of the unique isoetid plant communities. We tested whether the presence or disappearance of the isoetid Littorella uniflora (L.) could be predicted from environmental parameters, soil types and land use in the catchment area, and atmospheric nitrogen deposition. 2. We found that the topographic catchment area of a lake was an irrelevant unit to study effects of soil type and land use. Instead, using a GIS‐generated buffer zone around the lakes it proved feasible to classify 472 lakes into historical (if L. uniflora had disappeared) or recent (if L. uniflora was still present) Littorella lakes, based on soil type and land use. Our analysis showed that aeolian sand deposits and heath in the buffer zone favoured the presence of L. uniflora, whereas moraine clay and agriculture were strongly linked to the disappearance of L. uniflora. 3. However, in order to understand fully the presence or disappearance of L. uniflora, environmental data were needed in addition to soil types, land use and nitrogen deposition, and the use of discriminant analysis allowed us to classify 96% of the investigated lakes correctly into recent or historical sites. Alkalinity, total phosphorus, total nitrogen, aeolian sand deposits and heath were the most important parameters explaining the presence or disappearance of L. uniflora. Our analysis also indicated that eutrophication, rather than acidification, has likely caused the disappearance of L. uniflora from 218 of the 472 lakes investigated. 4. Our findings have widespread implications for the conservation or restoration of isoetid habitats and we recommend applying a wide buffer zone around lakes, with restrictions on farming and traditional forestry activities. In addition, our buffering concept may prove a useful tool for aquatic ecologists to investigate relationships between catchment features and organisms (plants, insects and amphibians) with aquatic as well as terrestrial life forms.     

Where temperate meets tropical: multi-factorial effects of elevated CO2, nitrogen enrichment, and competition on a mangrove-salt marsh community

Our understanding of how elevated CO₂ and interactions with other factors will affect coastal plant communities is limited. Such information is particularly needed for transitional communities where major vegetation types converge. Tropical mangroves (Avicennia germinans) intergrade with temperate salt marshes (Spartina alterniflora) in the northern Gulf of Mexico, and this transitional community represents an important experimental system to test hypotheses about global change impacts on critical ecosystems. We examined the responses of A. germinans (C₃) and S. alterniflora (C₄), grown in monoculture and mixture in mesocosms for 18 months, to interactive effects of atmospheric CO₂ and pore water nitrogen (N) concentrations typical of these marshes. A. germinans, grown without competition from S. alterniflora, increased final biomass (35%) under elevated CO₂ treatment and higher N availability. Growth of A. germinans was severely curtailed, however, when grown in mixture with S. alterniflora, and enrichment with CO₂ and N could not reverse this growth suppression. A field experiment using mangrove seedlings produced by CO₂‐ and N‐enriched trees confirmed that competition from S. alterniflora suppressed growth under natural conditions and further showed that herbivory greatly reduced survival of all seedlings. Thus, mangroves will not supplant marsh vegetation due to elevated CO₂ alone, but instead will require changes in climate, environmental stress, or disturbance to alter the competitive balance between these species. However, where competition and herbivory are low, elevated CO₂ may accelerate mangrove transition from the seedling to sapling stage and also increase above‐ and belowground production of existing mangrove stands, particularly in combination with higher soil N.     

Production and nutrient dynamics of plant communities on a sub-Antarctic island

Summary Studies of plant standing crop and nutrient concentrations have enabled an assessment of the seasonal changes in nutrient standing stocks (the mass of nutrients per m²) in two mire-grasslands at Marion Island (46°54S, 37°45E). Mire-grasslands are an important component of the island's vegetation, occurring on very wet peats and dominated by graminoids and bryophytes. Peak aboveground standing stocks of N, P and K in the vascular plant species of the mire-grasslands mostly occurred earlier in the season than did peak aboveground biomass, implying that aboveground accumulation rates of these nutrients were greater than the rate of biomass accumulation. Maximum Ca standing stocks coincided in the season with peak shoot biomass. Depending on the plant species, peak Mg and Na standing stocks occurred either before, or later than, peak shoot biomass. Total (above-plus belowground) standing stocks of nutrients (N+P+K+Ca+Mg+Na) at the time of peak aboveground biomass were 51 g m^-2 at study mire 1 and 44 g m^-2 at study mire 2. The most abundant element in the vegetation was N, followed by K. The net quantities of most nutrients translocated into the aboveground growth were mostly greater than the seasonal mean standing stocks in the aerial biomass. Except for Ca, nutrient standing stocks in the vegetation of the mire-grasslands are in the upper part of the range reported for sub-Arctic and Arctic graminoid communities. They are more similar to standing stocks at oceanic moorlands, montane grasslands and heath communities. Low Ca concentrations occur in the plants so that Ca standing stocks are lower than in most comparable northern hemisphere communities. Pool sizes (i.e. total quantities contained in the plant/soil system to a depth of 25 cm) of N, P, K and Ca are in the lower part of the range reported for wet, graminoid-dominated tundra and tundra-like communities of the northern hemisphere.     

Stability of above-ground and below-ground processes to extreme drought in model grassland ecosystems: Interactions with plant species diversity and soil nitrogen availability

Extreme drought events have the potential to cause dramatic changes in ecosystem structure and function, but the controls upon ecosystem stability to drought remain poorly understood. Here we used model systems of two commonly occurring, temperate grassland communities to investigate the short-term interactive effects of a simulated 100-year summer drought event, soil nitrogen (N) availability and plant species diversity (low/high) on key ecosystem processes related to carbon (C) and N cycling. Whole ecosystem CO₂ fluxes and leaching losses were recorded during drought and post-rewetting. Litter decomposition and C/N stocks in vegetation, soil and soil microbes were assessed 4 weeks after the end of drought. Experimental drought caused strong reductions in ecosystem respiration and net ecosystem CO₂ exchange, but ecosystem fluxes recovered rapidly following rewetting irrespective of N and species diversity. As expected, root C stocks and litter decomposition were adversely affected by drought across all N and plant diversity treatments. In contrast, drought increased soil water retention, organic nutrient leaching losses and soil fertility. Drought responses of above-ground vegetation C stocks varied depending on plant diversity, with greater stability of above-ground vegetation C to drought in the high versus low diversity treatment. This positive effect of high plant diversity on above-ground vegetation C stability coincided with a decrease in the stability of microbial biomass C. Unlike species diversity, soil N availability had limited effects on the stability of ecosystem processes to extreme drought. Overall, our findings indicate that extreme drought events promote post-drought soil nutrient retention and soil fertility, with cascading effects on ecosystem C fixation rates. Data on above-ground ecosystem processes underline the importance of species diversity for grassland function in a changing environment. Furthermore, our results suggest that plant–soil interactions play a key role for the short-term stability of above-ground vegetation C storage to extreme drought events.     

Alkalinity generation and sediment CO2 uptake influence establishment of Sparganium angustifolium in softwater lakes

1. Softwater lakes are generally dominated by slow growing, small, isoetid plant species that are adapted to the carbon‐ and nutrient‐limited conditions in these lakes. We investigated the strategy of a fast growing species, Sparganium angustifolium, for occupying softwater lakes. A field survey was carried out in Norwegian carbon‐limited Isoëteto‐Lobelietum softwater lakes to compare abiotic conditions at locations with and without S. angustifolium. In addition, long term abiotic changes (1995–2008) related to the sudden establishment of the species on experimentally limed plots were studied. Based on the results, the carbon acquisition mechanism of S. angustifolium was tested in eco‐physiological laboratory experiments. 2. The redox potential was significantly lower at locations with S. angustifolium (220 ± 2.3) compared to locations without S. angustifolium (338.1 ± 13.9). The lower redox potential was accompanied by significantly higher concentrations of HCO₃^?, CO₂ and Fe²⁺ in the sediment pore water, indicating in‐lake alkalinity generation due to higher iron reduction rates in the generally iron‐rich sediments. In addition, the lower redox potential was accompanied by a higher nutrient availability (NH₄⁺ and PO₄^3?) in the sediment pore water. Since there were no differences in water quality between the lakes, the ability of S. angustifolium to grow in softwater lakes very likely depends upon the higher dissolved inorganic carbon (DIC) and nutrient concentrations present in the sediment pore water. 3. Results from the liming experiment revealed that appearance of S. angustifolium on limed plots was related to the dissolution of Ca and Mg carbonates and development of a lower redox potential in the sediment. These processes were accompanied by a sustained increase in the availability of DIC in the sediment pore water. 4. The eco‐physiological experiments indicated that S. angustifolium can increase in biomass and produce floating leaves at a relatively high DIC availability in the root medium. In addition, it appeared that S. angustifolium can take up CO₂ by the roots. As far as we know, the ability to use sediment CO₂ has only been described as an adaptation typical for isoetid plant species. Use of the relatively large sediment CO₂ pools present in these sediment types (>1000 μmol L^?1) to enable development of long floating leaves for additional uptake of atmospheric CO₂ is a very different strategy to colonise softwater lakes as compared to isoetid plant species.     

Utilization of Sediment CO2 by Selected North American Isoetids

The photosynthetic uptake of root-zone CO₂ was determined forEriocaulon septangulare, Gratiola aurea, Isoetes macrospora,Littorella uniflora var. americana and Lobelia dortmanna aspart of a study of the photosynthetic carbon economy of submergedaquatic isoetids. The pH and dissolved inorganic carbon (DIC)of the sediment interstitial water in four Wisconsin lakes reflectedthe water column character, where the DIC increased with depthin the sediment to concentrations five to ten times those ofthe water column. Sediment free CO₂ concentrations were 5–50times those in the water column and were similar at all sites(about 05–1.0mM CO₂ in the root-zone). In ‘pH-drift’studies these plants were unable to take up HCO₂^–. Laboratory determinations of the carbon uptake from the rootand shoot-zones were made for all five species. These experimentsshowed that CO₂ in the root-zone accounted for 65–95 percent of external carbon uptake for the five species. For G.aurea and E. septangulare, root-zone CO₂ was > 85 per centof carbon uptake. Carbon, CO₂, photosynthesis, sediment, isoetid, Eriocaulon septangulare, Gratiola aurea, Isoetes macrospora, Littorella uniflora, Lobelia dortmanna     

Rapid oxygen exchange across the leaves of Littorella uniflora provides tolerance to sediment anoxia

1. Littorella uniflora and Lobelia dortmanna are prominent small rosette species in nutrient‐poor, soft‐water lakes because of efficient root exchange of CO₂ and O₂. We hypothesise that higher gas exchange across the leaves of L. uniflora than of L. dortmanna ensures O₂ uptake from water and underlies its greater tolerance to sediment anoxia following organic enrichment. 2. We studied plant response to varying sediment O₂ demand and biogeochemistry by measuring photosynthesis, gas exchange across leaves and O₂ dynamics in plants during long‐term laboratory and field studies. Frequent non‐destructive sampling of sediment pore water was used to track changes in sediment biogeochemistry. 3. Addition of organic matter triggered O₂ depletion and accumulation of , Fe²⁺ and CO₂ in sediments. Gas exchange across leaf surfaces was 13–16 times higher for L. uniflora than for L. dortmanna. Oxygen in the leaf lacunae of L. uniflora remained above 10 kPa late at night on anoxic sediments despite organic enrichment. Leaf content of N and P of L. uniflora remained sufficient to keep up photosynthesis despite prolonged sediment anoxia, whereas nutrient content was too low for long‐term survival of L. dortmanna. 4. High gas exchange across L. uniflora leaves improves its performance and survival on anoxic sediments compared with L. dortmanna. Lobelia dortmanna uses the same gas‐tight leaves in air and water, which makes it highly susceptible to sediment anoxia but more cost‐effective in ultra‐oligotrophic environments because of slow leaf turnover.