Sphagnum bleaching: Bicarbonate 'toxicity' and tolerance for seven Sphagnum species.

Growth and functioning of Sphagnum mosses are closely linked to water level and chemistry. Sphagnum mosses occur in wet, generally acidic conditions, and when buffered, alkaline water is known to negatively impact Sphagnum. The effects of time, dose and species-specific responses of buffered, alkaline water on Sphagnum are largely unknown. We investigated the effects of bicarbonate and calcium on the survival, growth and physiological functioning of seven Sphagnum species occurring in contrasting environments, from raised bogs to (rich) fens. Mosses were submerged in different concentrations of bicarbonate and calcium solutions for 10 weeks under climate-controlled circumstances. After 2 weeks, all species exposed to the high bicarbonate treatment (2.0 mM) showed severe potassium leakage and swift discoloration. In contrast, species showed differential responses to the intermediate bicarbonate treatment (0.8 mM), some with a later onset of potassium leakage. S. squarrosum, S. teres & S. contortum generally persisted the longest, with all species dying after 6 to 10 weeks. Calcium alone, in contrast, negatively affected S. squarrosum, S. teres & S. contortum, causing discoloration and potassium leakage. Our study shows enrichment with bicarbonate, but not calcium, is detrimental for most Sphagnum species tested. A mechanistic model was developed that is consistent with dose and duration dependence and the species specificity. Future conservation and restoration measures for Sphagnum-dominated habitats and Sphagnum farming (cultivation, production and harvest of Sphagnum moss biomass) should limit flooding with bicarbonate-rich waters while investigating new management options, like acidifying surface waters to lower bicarbonate levels.

Differential responses of two wetland graminoids to high ammonium at different pH values.

Enhanced soil ammonium (NH4+) concentrations in wetlands often lead to graminoid dominance, but species composition is highly variable. Although NH4+ is readily taken up as a nutrient, several wetland species are known to be sensitive to high NH4+ concentrations or even suffer toxicity, particularly at low soil pH. More knowledge about differential graminoid responses to high NH4+ availability in relation to soil pH can help to better understand vegetation changes. The responses of two wetland graminoids, Juncus acutiflorus and Carex disticha, to high (2 mmol·l(-1) ) versus control (20 µmol·l(-1) ) NH4+ concentrations were tested in a controlled hydroponic set up, at two pH values (4 and 6). A high NH4+ concentration did not change total biomass for these species at either pH, but increased C allocation to shoots and increased P uptake, leading to K and Ca limitation, depending on pH treatment. More than 50% of N taken up by C. disticha was invested in N-rich amino acids with decreasing C:N ratio, but only 10% for J. acutiflorus. Although both species appeared to be well adapted to high NH4+ loadings in the short term, C. disticha showed higher classic detoxifying responses that are early warning indicators for decreased tolerance in the long term. In general, the efficient aboveground biomass allocation, P uptake and N detoxification explain the competitive strength of wetland graminoids at the expense of overall biodiversity at high NH4+ loading. In addition, differential responses to enhanced NH4+ affect interspecific competition among graminoids and lead to a shift in vegetation composition.

Massive uprooting of Littorella uniflora (L.) Asch. during a storm event and its relation to sediment and plant characteristics.

During spring storms massive uprooting of Littorella uniflora occurred in a shallow Dutch softwater lake. The aim of this study was to test whether changes in plant morphology and sediment characteristics could explain the observed phenomenon. Uprooting was expected to occur in plants having a high shoot biomass and low root to shoot ratio (R:S), growing on sediments with a high organic matter content. Normally, uprooting of the relative buoyant L. uniflora is prevented by an extensive root system, expressed as a high R:S. This was studied by sampling floating and still rooted L. uniflora plants, as well as sediment and sediment pore water, along a gradient of increasing sediment organic matter content. Increasing organic matter content was related to increasing L. uniflora shoot biomass and consequently decreasing R:S. Furthermore, the results indicated that uprooting indeed occurred in plants growing on very organic sediments and was related to a low R:S. The increased shoot biomass on more organic sediments could be related to increased sediment pore water total inorganic carbon (TIC; mainly CO2 ) availability. Additionally, increased phosphorus availability could also have played a role. The disappearance of L. uniflora might lead to higher nutrient availability in the sediments. It is suggested that this could eventually promote the expansion of faster-growing macrophytes.

Nutrient additions in pristine Patagonian Sphagnum bog vegetation: can phosphorus addition alleviate (the effects of) increased nitrogen loads.

Sphagnum-bog ecosystems have a limited capability to retain carbon and nutrients when subjected to increased nitrogen (N) deposition. Although it has been proposed that phosphorus (P) can dilute negative effects of nitrogen by increasing biomass production of Sphagnum mosses, it is still unclear whether P-addition can alleviate physiological N-stress in Sphagnum plants. A 3-year fertilisation experiment was conducted in lawns of a pristine Sphagnum magellanicum bog in Patagonia, where competing vascular plants were practically absent. Background wet deposition of nitrogen was low (≈ 0.1-0.2 g · N · m(-2) · year(-1)). Nitrogen (4 g · N · m(-2) · year(-1)) and phosphorus (1 g · P · m(-2) · year(-1)) were applied, separately and in combination, six times during the growing season. P-addition substantially increased biomass production of Sphagnum. Nitrogen and phosphorus changed the morphology of Sphagnum mosses by enhancing height increment, but lowering moss stem density. In contrast to expectations, phosphorus failed to alleviate physiological stress imposed by excess nitrogen (e.g. amino acid accumulation, N-saturation and decline in photosynthetic rates). We conclude that despite improving growth conditions by P-addition, Sphagnum-bog ecosystems remain highly susceptible to nitrogen additions. Increased susceptibility to desiccation by nutrients may even worsen the negative effects of excess nitrogen especially in windy climates like in Patagonia.

Selecting predictors for discriminant analysis of species performance: an example from an amphibious softwater plant.

Selecting an appropriate variable subset in linear multivariate methods is an important methodological issue for ecologists. Interest often exists in obtaining general predictive capacity or in finding causal inferences from predictor variables. Because of a lack of solid knowledge on a studied phenomenon, scientists explore predictor variables in order to find the most meaningful (i.e. discriminating) ones. As an example, we modelled the response of the amphibious softwater plant Eleocharis multicaulis using canonical discriminant function analysis. We asked how variables can be selected through comparison of several methods: univariate Pearson chi-square screening, principal components analysis (PCA) and step-wise analysis, as well as combinations of some methods. We expected PCA to perform best. The selected methods were evaluated through fit and stability of the resulting discriminant functions and through correlations between these functions and the predictor variables. The chi-square subset, at P < 0.05, followed by a step-wise sub-selection, gave the best results. In contrast to expectations, PCA performed poorly, as so did step-wise analysis. The different chi-square subset methods all yielded ecologically meaningful variables, while probable noise variables were also selected by PCA and step-wise analysis. We advise against the simple use of PCA or step-wise discriminant analysis to obtain an ecologically meaningful variable subset; the former because it does not take into account the response variable, the latter because noise variables are likely to be selected. We suggest that univariate screening techniques are a worthwhile alternative for variable selection in ecology.

The Eocene Arctic Azolla bloom: environmental conditions, productivity and carbon drawdown.

Enormous quantities of the free-floating freshwater fern Azolla grew and reproduced in situ in the Arctic Ocean during the middle Eocene, as was demonstrated by microscopic analysis of microlaminated sediments recovered from the Lomonosov Ridge during Integrated Ocean Drilling Program (IODP) Expedition 302. The timing of the Azolla phase (approximately 48.5 Ma) coincides with the earliest signs of onset of the transition from a greenhouse towards the modern icehouse Earth. The sustained growth of Azolla, currently ranking among the fastest growing plants on Earth, in a major anoxic oceanic basin may have contributed to decreasing atmospheric pCO2 levels via burial of Azolla-derived organic matter. The consequences of these enormous Azolla blooms for regional and global nutrient and carbon cycles are still largely unknown. Cultivation experiments have been set up to investigate the influence of elevated pCO2 on Azolla growth, showing a marked increase in Azolla productivity under elevated (760 and 1910 ppm) pCO2 conditions. The combined results of organic carbon, sulphur, nitrogen content and 15N and 13C measurements of sediments from the Azolla interval illustrate the potential contribution of nitrogen fixation in a euxinic stratified Eocene Arctic. Flux calculations were used to quantitatively reconstruct the potential storage of carbon (0.9-3.5 10(18) gC) in the Arctic during the Azolla interval. It is estimated that storing 0.9 10(18) to 3.5 10(18) g carbon would result in a 55 to 470 ppm drawdown of pCO2 under Eocene conditions, indicating that the Arctic Azolla blooms may have had a significant effect on global atmospheric pCO2 levels through enhanced burial of organic matter.

Toxicity of reduced nitrogen in eelgrass (Zostera marina) is highly dependent on shoot density and pH.

In sheltered, eutrophicated estuaries, reduced nitrogen (NHx), and pH levels in the water layer can be greatly enhanced. In laboratory experiments, we studied the interactive effects of NHx, pH, and shoot density on the physiology and survival of eelgrass (Zostera marina). We tested long-term tolerance to NHx at pH 8 in a 5-week experiment. Short-term tolerance was tested for two shoot densities at both pH 8 and 9 in a 5-day experiment. At pH 8, eelgrass accumulated nitrogen as free amino acids when exposed to high loads of NHx, but showed no signs of necrosis. Low shoot density treatments became necrotic within days when exposed to NHx at pH 9. Increased NH3 intrusion and carbon limitation seemed to be the cause of this, as intracellular NHx could no longer be assimilated. Remarkably, experiments with high shoot densities at pH 9 showed hardly any necrosis, as the plants seemed to be able to alleviate the toxic effects of high NHx loads through joint NHx uptake. Our results suggest that NHx toxicity can be important in worldwide observed seagrass mass mortalities. We argue that the mitigating effect of high seagrass biomass on NHx toxicity is a positive feedback mechanism, potentially leading to alternative stable states in field conditions.

Effects of mining activities on heavy metal concentrations in water, sediment, and macroinvertebrates in different reaches of the Pilcomayo River, South America.

From 1997 until 1999 the extent and the ecological effects of zinc, copper, lead, and cadmium pollution were studied in different reaches of the South American Pilcomayo River. A comparison of metal concentrations in water, sediment, and chironomid larvae, as well as the diversity of macroinvertebrate species, was made between sites near the origin of the Pilcomayo River, with hardly any mining activities, sites in the Potosí region, with intensive mining, and sites located 500 km or further downstream of Potosí, in the Chaco plain. Samples were also collected in an unpolluted river (Cachi Mayu River) and in the Tarapaya River, which is strongly contaminated by mine tailings (1000 tons a day). The upper parts of the Pilcomayo River are strongly affected by the release of mine tailings from the Potosí mines where mean concentrations of lead, cadmium, copper, and zinc in water, filtered water, sediment, and chironomid larvae were up to a thousand times higher than the local background levels. The diversity of the benthic macroinvertebrate community was strongly reduced in the contaminated parts; 97% of the benthic macroinvertebrates consisted of chironomid larvae. The degree of contamination in the lower reaches of the river, however, was fairly low because of sedimentation processes and the strong dilution of mine tailings with enormous amounts of clean sediment from erosion processes. Analysis of sediment cores from the Ibibobo floodplain, however, reveal an increase of the heavy metal concentrations in the lower reaches since the introduction of the contaminating flotation process in the mine industry in 1985.

Potential sensitivity of mires to drought, acidification and mobilisation of heavy metals: the sediment s/(Ca + Mg) ratio as diagnostic tool.

In recent decades sulphate concentrations in the ground water in many parts of The Netherlands have increased dramatically resulting in increased production of iron-(di)sulphides in sediments of ecosystems fed by this water. A sediment survey was carried out to study the potential sensitivity of wetlands to drought and subsequent acidification as a consequence of iron-(di)sulphide oxidation. Dessication led to severe acidification and mobilisation of heavy metals when the sediment S/(Ca + Mg) ratio exceeded 2/ 3. A total of 47% of the investigated locations contained S/(Ca + Mg) ratios higher than 2/3 and in 100, 75 and 50% of the locations mobilisation of Zn, Cd and Ni exceeded the Dutch signal value for ground water. Consistent with the sediment survey, lime addition experiments confirmed that increasing the buffer capacity, down to a S/(Ca + Mg) ratio 2/3, led to a drastic inhibition of the acidification and heavy metal percolation from dredged sediments. The performance of the same processes under drained field conditions demonstrates the relevance of these processes during dry summers.

The roles of internal iron hydroxide precipitation, sulphide toxicity and oxidizing ability in the survival of Stratiotes aloides roots at different iron concentrations in sediment pore water.

Stratiotes aloides L. is an aquatic macrophyte that occurs in waters on reduced peaty sediments which have a relatively narrow range of free-iron content in the sediment. Comparison of different aquatic macrophytes reveals that species from reducing sediments have much lower oxidizing ability than do species from oxidizing sediments. Compared with those other species from reducing sediments, the oxidizing ability of Stratiotes aloides is very low and probably makes the species very vulnerable to sulphide toxicity and internal precipitation of iron hydroxide. Apoplastic iron contents were determined for Stratiotes roots growing in sediments with different free-iron concentrations. Roots collected from sediment with a low free-iron content appeared to have a relatively low apoplastic-iron content whereas roots from sediments with a relatively high free-iron concentration had a relatively high apoplastic-iron content. In sediments with low free-iron levels, sulphide levels are generally high. Using light microscopy, iron hydroxide precipitates were observed around the endodermis and cortical air spaces of the roots of Stratiotes aloides growing in iron-rich sediments. Internal iron oxidation outside the endodermis can prevent iron toxicity inside the stele and thus enable root apices to survive. The root hairs, however, die because of iron hydroxide precipitation at their base; the dead roots have the highest visible iron hydroxide content. Excessive internal iron hydroxide precipitation and the consequential early death of the roots probably explains the absence of Stratiotes aloides in locations with high free-iron levels in sediment pore water.