Impact of acidification and eutrophication on macrophyte communities in soft waters. II. Experimental studies

In The Netherlands, there has been a dramatic decline during the last 30 years in the number of stands belonging to the phytosociological alliance Littorellion. Generally, the communities classified within this alliance occur in poorly buffered, oligotrophic waters, with very low phosphate, nitrogen and carbon dioxide levels in the water layer and considerably higher nutrient levels in the sediment. The plant species dominating these communities are isoetids such as Litoorella uniflora (L.) Aschers., Lobelia dortmanna L. and Isoetes lacustris L., which show various adaptations to make successful growth possible under these conditions.Field observations showed that the water where Littorella uniflora had disappeared or strongly decreased could be divided into two groups. A major group (77%) was characterized by the presence of submerged Juncus bulbosus L. and/or Sphagnum species. These water appeared to be strongly acidified (pH < 4.5) and had increased nitrogen levels with ammonium as the dominant N-source. Within this group, the waters with luxuriant growth of Juncus bulbosus and/or Sphagnum spp. had strongly increased carbon dioxide levels in both sediment and water.Different types of experiments proved causal relationships between the observed changes in macrophytes and the changed physico-chemical parameters. Ecophysiological experiments showed that Juncus bulbosus lacks the typical adaptations of the isoetid plant species, i.e. it uses very low amounts of sediment-CO₂ and releases only a little oxygen from the roots. However, Juncus bulbosus is more able than Littorella uniflora to use CO₂ from the water layer. From the nutrient-uptake experiments, the decreased nitrate and increased ammonium levels seem to be favourable to Juncus bulbosus. The culture experiments clearly demonstrated that the biomass of Juncus bulbosus only increased strongly when the sediment was poorly buffered and the pH of water was low. When combining factors like CO₂ enrichment of the sediment, with and without phosphate, and/or ammonium enrichment of the water in the culture experiments, it is clearly shown that phosphate and/or ammonium enrichment without CO₂ enrichment do not lead to an increase in biomass of Juncus bulbosus. Therefore, it is obvious that the changes in the macrophyte community can be ascribed primarily to changes in the carbon budget as a result of acidification.A minor group of waters (23%) was characterized by the absence of submerged Juncus bulbosus and/or Sphagnum spp. In most of these waters, submerged plant species occurred, such as Myriophyllum alterniflorum DC or non-rooted species such as Riccia fluitans L. These waters were not acidified, and generally had an increased alkalinity and higher nitrogen and phosphate levels of sediment and/or water. Culture experiments showed that phosphate enrichment of the sediment alone leads to luxuriant growth of submerged macrophyte species such as Myriophyllum alterniflorum, whereas phosphate enrichment of both sediment and water leads to mass development of non-rooted plant species such as Riccia fluitans.     

Impact of acidification and eutrophication on macrophyte communities in soft waters in The Netherlands I. Field observations

During the last decades a strong decline has been noticed in the number of waters dominated by “Littorellion” species, mostly isoetids such as Lobelia dortmanna L., Isoetes lacustris L. and Littorella uniflora (L.) Aschers. Sixty-eight waters, which were known to be dominated by L. uniflora after 1950 were investigated. In 1980, L. uniflora appeared to be absent or to have strongly decreased in 53 (78%) of the waters. In 41 of them, Littorella had been replaced by submerged Juncus bulbosus L. and/or Sphagnum spp. These changes seem to have been caused by changed inorganic carbon budgets as a consequence of acidification.In the remaining 12 waters, eutrophication of the water and/or sediment seems to be responsible for the changes in the plant communities. Enrichment with phosphate of the mineral sediment alone, leads to luxurious growth of submerged, rooted macrophyte species such as Myriophyllum alterniflorum DC and Ranunculus peltatus Schrank, whereas phosphate-enrichment of both sediment and water leads to luxurious growth of pleustophytes such as Riccia fluitans L. and Lemna minor L. in small, shallow waters, and to plankton bloom and luxurious growth of epiphytes in larger, deeper waters.In these cases light limitation seems to be responsible for the disappearance or decline of the “Littorellion” species.     

Effect of Lake Trophic Status and Rooted Macrophytes on Community Composition and Abundance of Ammonia-Oxidizing Prokaryotes in Freshwater Sediments

Communities of ammonia-oxidizing archaea (AOA) and bacteria (AOB) in freshwater sediments and those in association with the root system of the macrophyte species Littorella uniflora, Juncus bulbosus, and Myriophyllum alterniflorum were compared for seven oligotrophic to mesotrophic softwater lakes and acidic heathland pools. Archaeal and bacterial ammonia monooxygenase alpha-subunit (amoA) gene diversity increased from oligotrophic to mesotrophic sites; the number of detected operational taxonomic units was positively correlated to ammonia availability and pH and negatively correlated to sediment C/N ratios. AOA communities could be grouped according to lake trophic status and pH; plant species-specific communities were not detected, and no grouping was apparent for AOB communities. Relative abundance, determined by quantitative PCR targeting amoA, was always low for AOB (<0.05% of all prokaryotes) and slightly higher for AOA in unvegetated sediment and AOA in association with M. alterniflorum (0.01 to 2%), while AOA accounted for up to 5% in the rhizospheres of L. uniflora and J. bulbosus. These results indicate that (i) AOA are at least as numerous as AOB in freshwater sediments, (ii) aquatic macrophytes with substantial release of oxygen and organic carbon into their rhizospheres, like L. uniflora and J. bulbosus, increase AOA abundance; and (iii) AOA community composition is generally determined by lake trophy, not by plant species-specific interactions.Oxygen release from the roots of macrophyte species such as Littorella uniflora (L.) Asch. (shore weed), Lobelia dortmanna L. (water lobelia), and Glyceria maxima (Hartm.) Holmb. (reed sweet grass) stimulates nitrification and coupled nitrification-denitrification in the rhizosphere compared to that in unvegetated sediment (2, 36, 40). These interactions are of high ecological relevance especially in oligotrophic systems, since enhanced nitrogen loss due to rhizosphere-associated denitrification can retard natural eutrophication and succession of plant communities (1). While the microbial communities involved in coupled nitrification-denitrification have been well studied in rice paddy soils (7, 11), less information is available for natural freshwater sediments, especially those from oligotrophic lakes (2, 26).The first key step of coupled nitrification-denitrification, the oxidation of ammonia to nitrite, is catalyzed by two groups of prokaryotes—the ammonia-oxidizing bacteria (AOB) (24) and the only recently recognized ammonia-oxidizing archaea (AOA) (22). For both groups, the gene encoding the alpha-subunit of ammonia monooxygenase (amoA) has been widely used as a functional marker to analyze their community compositions (15, 25); recent studies demonstrated the ubiquity of AOA and their predominance over AOB in a broad range of environments (32, 38). AOA, but not AOB, were also strongly enriched in the rhizosphere of the freshwater macrophyte Littorella uniflora in a mesotrophic Danish lake, suggesting that AOA were primarily responsible for increased rates of nitrification in the rhizosphere of this plant species (19). Moreover, ammonia oxidizer communities differed between rhizosphere and unvegetated sediment, indicating a plant-specific effect on AOA and AOB community composition. The objectives of this study were therefore to test whether (i) AOA generally predominate over AOB in freshwater sediments and especially in macrophyte rhizospheres and (ii) macrophytes have species-specific effects on abundance and community composition of AOA and AOB in rhizosphere sediments and on root surfaces.To address these questions, two shallow heathland pools and five lakes in Denmark and Germany, ranging from low-pH and dystrophic sites to neutral-pH and oligotrophic and mesotrophic sites, were chosen, and three macrophyte species—Littorella uniflora, Juncus bulbosus L. (bulbous rush), and Myriophyllum alterniflorum DC. (alternate water milfoil)—were selected as model systems. These plant species differ in nitrogen nutrition, extent of radial oxygen loss, and lifestyle, presumably resulting in differential, plant species-specific effects on rhizosphere- and root-associated AOA and AOB communities. L. uniflora prefers nitrate as the nitrogen source, while J. bulbosus prefers ammonium (41, 45); oxygen release is high to moderate from the roots of L. uniflora and J. bulbosus (9, 12) but is minor from the roots of M. alterniflorum (M. Herrmann, P. Stief, and A. Schramm, unpublished results); L. uniflora and J. bulbosus remain photosynthetically active throughout the year, while only the below-ground parts of M. alterniflorum are retained during winter.Rhizosphere sediments and roots from each plant species were sampled from three different sites per species, and unvegetated sediment was obtained from all seven sites. The comparison of samples from these different sites and compartments (rhizosphere, root surface, unvegetated sediment) allowed an evaluation of the importance of plant species relative to that of environmental conditions related to lake trophic status and pH on ammonia oxidizer communities.     

The effects of amino acids and ammonium on the growth of plant cells in suspension culture

A suspension culture of soybean (Glycine max L.) was grown on a defined medium in which the nitrogen sources were nitrate (25 mM) and ammonium (2 mM). The cells did not grow on nitrate unless the medium was supplemented with ammonium or glutamine. The l- and d-isomers of 12 amino acids tested singly could not replace ammonium. Most amino acids (4 mM) inhibited growth when the cells were cultured on nitrate and ammonium. Cells from five other plants (Reseda luteoli L.; Triticum monococcum L.; flax, Linum usitatissimum L.; horseradish, Amoracia lapathifolia Gilib; Haplopappus gracilis L.) grew on the defined medium with nitrate (25 mM) as the sole nitrogen source. Higher cell yields were obtained when ammonium (2 mM) or glutamine also was present. Supplementing the defined medium with high concentrations of ammonium (20 mM) inhibited growth of soybean, Haplopappus, and wheat cells. Addition of citrate (5 mM) relieved the inhibitory effects of ammonium in soybean and wheat cells but not in the Haplopappus cells.     

A model on the competition between two macrophyte species in acidifying shallow soft-water lakes in the Netherlands

To predict changes in vegetation as a result of acidification in shallow Dutch lakes a mathematical model was formulated which describes the growth ofLittorella uniflora andJuncus bulbosus, species characteristic for non-acidified and acidified situations, respectively. The two species compete for light, CO₂ and nutrients. The model integrates current knowledge about the ecophysiology of the species. The results generated by the model are similair to field data concerning dominance of the species in different habitats.L. uniflora is able to grow well under environmental conditions associated with acidification, as long as it is not outcompeted byJ. bulbosus. The competitive advantage of morphological adaptions of the macrophyte species are discussed in relation to the model results.     

Comparison of nitrogen uptake in the roots and rhizomes of <Emphasis Type="Italic">Leymus chinensis</Emphasis>

Leymus chinensis (Trin.) Tzvel is a rhizomatous grass species in the Eastern Eurasian steppe zone that is often limited by low soil nitrogen availability. Although a previous study showed that the rhizomes of L. chinensis have the capacity to take up nitrogen, the importance of such uptake for nitrogen nutrition is unclear. Moreover, little is known regarding the inorganic nitrogen uptake kinetics of roots and rhizomes in response to nitrogen status. Here, we first found that ammonium is preferred over nitrate and glycine for L. chinensis growth. Using the ¹⁵N-labelling method, we found that the rate of ion influx into roots was approximately five-fold higher than into rhizomes under the same nitrogen content, and the ion influxes into roots and rhizomes under 0.05 mM N were greater than in the presence of 3 mM N, especially in the form of NH₄⁺. Using a non-invasive micro-test technique, we characterised the patterns of NH₄⁺ and NO₃^– fluxes in the root mature zone, root tip, rhizome mature zone, and rhizome tip following incubation in the solution with different N compounds and different N concentrations. These results suggest a dynamic balance between the uptake, utilisation, and excretion of nitrogen in L. chinensis.     

Parallelism in the evolution of rotifers

The carbon uptake by the roots of three common submersed macrophytes, Myriophyllum spicatum, Heteranthera dubia and Vallisneria americana, was measured in situ under a range of pH and dissolved inorganic carbon (DIC) concentrations. After 4–8 weeks of growth in ¹⁴C labelled sediments, less than 1.5% of the total C found in the shoots originated from root uptake. Between species, significant differences in C uptake by the root were found and root development alone could explain most of the observed variance (84%). C uptake by the roots of aquatic macrophytes thus appears to be restricted to small, rosette-shaped plants growing in poorly mineralized waters.     

Nuisance growth of Juncus bulbosus: the roles of genetics and environmental drivers tested in a large‐scale survey

1. Invasive species can transform aquatic ecosystems, and the nuisance growth of the freshwater macrophyte Juncus bulbosus has become a problem in many lakes and rivers in northern Europe. It affects biodiversity strongly and conflicts with human uses, not least compromising the generation of hydroelectricity. The causes of the proliferation of these massive stands of J. bulbosus are not finally resolved, however. 2. In this study, a wide range of catchment, lake and sediment parameters (n = 34) were assessed for 139 lakes in Southern Norway, with the aim of explaining the presence or absence of J. bulbosus and to assess potential drivers behind its prolific growth. 3. Juncus bulbosus was more often present in lakes with lower pH and phosphate concentrations, and a higher element ratio of dissolved inorganic nitrogen (DIN) to total phosphorus (DIN : TotP). 4. Despite the many parameters measured across substantial environmental gradients, none explained nuisance growth. Genetic screening (amplified fragment length polymorphism fingerprinting) of plants from a subset of lakes and additional river sites also showed no genetic differences between the various growth forms. A macrophyte trophic index, however, suggested that the most problematic growth occurred in the most oligotrophic lakes. 5. The lack of consistent patterns may reflect either factors not assessed in our survey, or that the current extension of stands represents a gradual cumulative response over time, not characterised effectively in our snapshot survey. Nevertheless, we can now exclude some putative causes of nuisance growth, including in particular genetics and N‐deposition.     

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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Elevated CO2 levels enhance the uptake and metabolism of organic nitrogen

The effects of elevated CO₂ (eCO₂) on the relative uptake of inorganic and organic nitrogen (N) are unclear. The uptake of different N sources by pak choi (Brassica chinensis L.) seedlings supplied with a mixture of nitrate, glycine and ammonium was studied using ¹⁵N‐labelling under ambient CO₂ (aCO₂) (350 ppm) or eCO₂ (650 ppm) conditions. ¹⁵N‐labelled short‐term uptake and ¹⁵N‐gas chromatography mass spectrometry (GC–MS) were applied to measure the effects of eCO₂ on glycine uptake and metabolism. Elevated CO₂ increased the shoot biomass by 36% over 15 days, but had little effect on root growth. Over the same period, the N concentrations of shoots and roots were decreased by 30 and 2%, respectively. Elevated CO₂ enhanced the uptake and N contribution of glycine, which accounted for 38–44% and 21–40% of total N uptake in roots and shoots, respectively, while the uptake of nitrate and ammonium was reduced. The increased glycine uptake resulted from the enhanced active uptake and enhanced metabolism in the roots. We conclude that eCO₂ may increase the uptake and contribution of organic N forms to total plant N nutrition. Our findings provide new insights into plant N regulation under eCO₂ conditions.     

Nitrogen acquisition in four co-existing species from an upland acidic grassland

The concentration of both nitrate and ammonium nitrogen was measured in soil taken from an upland acidic (pH 4.5) grassland habitat, containing four co-existing species, Deschampsia flexuosa (L.) Trin., Festuca ovina L., Juncus squarrosus L. and Nardus stricta L. Both nitrate and ammonium nitrogen were found to be present in the soil, in similarly small quantities. The effect of both sources of nitrogen on relative growth rate was studied, and an attempt was made to determine whether nitrate or ammonium nitrogen is the immediate source of nitrogen for these plants using assays of nitrate reductase (EC 1.6.6.2) and ammonium uptake. All four species showed larger growth rates on the same concentration of ammonium nitrogen compared to nitrate nitrogen. All species showed low activities of leaf nitrate reductase, even in plants grown on 18 mol nitrate m⁻³. Ammonium uptake activity appeared to be higher in species which showed the lowest nitrate reductase activity and least response to increasing nitrate concentration in the growth medium.     

The response of interstitial ammonium in eelgrass (Zostera marina L.) beds to environmental perturbations

Natural and human-induced perturbations of eelgrass (Zostera marina L.) beds were used to examine the interaction between the sediment interstitial ammonium pool and nitrogen uptake by the plants. Eelgrass colonization of unvegetated areas was accompanied by a substantial decrease in the interstitial ammonium pool over a 4-yr period. The changes in interstitial ammonium and shoot density during colonization support an already determined relationship between shoot density and ammonium pool measurements. In field perturbation experiments, removing eelgrass leaves and sealing the sediment surface altered the flux of ammonium from the interstitial ammonium pool, and resulted in a rapid increase in interstitial ammonium concentrations. Measurements of ammonium accumulation under the various perturbation conditions and a control permitted calculation of the sediment ammonium flux. These estimates include uptake by eelgrass roots, regeneration in the root zone, and diffusion from the sediments. Nitrogen limitation was observed in some eelgrass beds.     

Age and growth,reproduction and diet of a sublittoral population of the rock goby Gobius paganellus (Teleostei,Gobiidae)

Colonisation of extremely acidic waters (pH 3) by aquatic angiosperms occurs widely, but is poorly documented. Unlike acid rain affected and other naturally acidic aquatic ecosystems, waters with pH 3 usually have a high conductivity, with high concentrations of SO₄ ^2- and often high concentrations of Fe³⁺, other heavy metal ions and Al³⁺. Where Fe³⁺ concentration is high, as in many mine waters, it provides a strong buffering system. In such waters, the biogeochemical Fe cycle exerts over water chemistry and the availability of nutrients and carbon for organisms. Biological activity is limited by low concentrations of phosphorus and inorganic carbon (DIC), which in this pH range is essentially all in the form of dissolved CO₂. A number of angiosperms grow in such waters including Phragmites australis, Typha spp. and Juncus bulbosus, though the last is the only one reported to grow totally submerged in waters with pH 3 . J. bulbosus occurs in many lignite mining lakes in Lusatia (north eastern Germany) with pH 3. The characteristics and possible survival strategies for this and other species are discussed.     

Photosynthesis of submersed macrophytes in acidified lakes II. Carbon limitation and utilization of benthic CO2 sources

Photosynthetic characteristics of carbon-dioxide limitations were analyzed for leaf tissue in a Cartesian-diver system, in which irradiance could be stringently controlled, and with whole plants in electrode macrosystems for submerged macrophytes (Juncus bulbosus L., Sphagnum auriculatum Schimp. var. inundatum (Russow) M. O. Hill) and other benthic moss and algae (Drepanocladus, Batrachospermum, and an algal mat) from acidified lakes. Light compensation points were extremely low for Juncus (1.5–6 μE m^?2 s^?1) and Sphagnum (3–10), and higher for shallow-inhabiting Batrachospermum (22–33). Leaf tissue, whole plants, and algal populations were rapidly limited by CO₂ availability under closed submersed, acidified conditions (pH 4–6).Controlled and in situ experiments were performed, in which the rooting tissue of Juncus bulbosus was partitioned from the leaves and the rates of photosynthetic carbon fixation of the foliage, utilizing dissolved inorganic ¹⁴C-carbon from the water, were analyzed under different conditions of CO₂ enhancement in the rhizosphere of the sediments. Results demonstrated that: (a) from 25 to 40% of the carbon fixed in the leaves can originate from the rhizosphere, diffuse to the leaves via internal gas lacunae, and be fixed photosynthetically; (b) photosynthetic utilization of CO₂ from the water surrounding the leaves is reduced markedly when the CO₂ concentration of the rhizosphere was increased by direct additions of CO₂, bacteria, or organic compounds (glucose, acetate) that stimulate bacterial growth. Shifts to predominance of submersed benthic primary producers with low light compensation points and adapted to acidified lakes are related in part to circumvention of carbon limitation in the water by utilization of enhanced CO₂ availability in the rhizosphere and at the sediment—water interface from bacterial degradation of organic matter, and in part to physiological mechanisms that conserve and recycle CO₂ of respiration and photorespiration.     

Differences in nitrate and ammonium uptake between Scots pine and European larch

In forest soils, ammonium is usually the predominant form of inorganic nitrogen. However, the capacity of trees to utilize both NO₃ ^- and NH₃ ⁺ may provide greater flexibility in responding to changes of nitrogen supply from the environment. Such capacity has been studied in seedlings of Scots pine (Pinus sylvestris L.) and European larch (Larix decidua Mill.) grown in the presence or absence of either nitrate or ammonium. Nitrate-induced plants showed a higher nitrate uptake rate than non-induced plants; this difference was almost negligible after 24 h of exposure to NO₃ ^-. Ammonium uptake in both species was consistently higher than that of nitrate, regardless of prior nitrogen provision. In both nutrient conditions, larch showed a more efficient transport system in comparison with Scots pine, with higher ammonium and nitrate uptake rates in both induced and non-induced plants. This was consistent also with the activity of nitrate reductase, measured in vivo in roots and leaves. This revised version was published online in June 2006 with corrections to the Cover Date.     

Activities of Carboxylation Enzymes in Freshwater Macrophytes

Fifteen species of freshwater macrophytes, mainly from cool,temperate waters, were assayed for ribulose bisphosphate carboxylase-oxygenase(RuBPCase) and phosphoenolpyruvate carboxylase (PEPCase) activities.In extracts from all the species RuBPCase was the most activecarboxylation enzyme, and the RuBPCase/PEPCase ratio was atleast 2·0, even for the submersed species Isoetes lacustrisL. and Littorella unifiora (L.) Aschers. which have been reportedto show Crassulacean Acid Metabolism (CAM) activity. The PEPCaseactivity in I.lacustris was lower than that found in some non-CAM-likespecies. In this respect, I.lacustris and L unifiora differfrom most terrestrial CAM plants. However, these two species,along with Potamogeton praelongus Wulf. and Juncus bulbosusvar.fluitans L., had the lowest RuBPCASE/PEPCase ratios, lowerthan found in terrestrial C₃ species; suggesting that the potentialfor substantial photosynthetic metabolism of C₄ acids existsin some temperate, submersed plants. In the three amphibiousspecies (Potamogeton polygonifolius Pourr., Mentha aquaticaL., and Hippuris vulgaris L.) examined, the aerial leaves exhibitedhigher RuBPCase activities than the submersed leaves. Key words: Ribulose bisphosphate carboxylase-oxygenase, phosphoenolpruvate carboxylase, freshwater macrophytes     

Differential regulation of nitrate reductase induction in roots and shoots of cotton plants

The induction of nitrate reductase activity in root tips of cotton (Gossypium hirsutum L.) was regulated by several amino acids and by ammonium. Glycine, glutamine, and asparagine strongly inhibited induction of activity by nitrate and also decreased growth of sterile-cultured roots on a nitrate medium. Methionine, serine, and alanine weakly inhibited induction, and 11 other amino acids had little or no effect. Ammonium also decreased induction in root tips, but was most effective only at pH 7 or higher. The optimum conditions for ammonium regulation of induction were identical to those for growth of sterile-cultured roots on ammonium as the sole nitrogen source. Aspartate and glutamate strongly stimulated induction, but several lines of evidence indicated that the mechanism of this response was different from that elicited by the other amino acids. The effects of amino acids on induction appeared to be independent of nitrate uptake.     

Successional changes in the soft-water macrophyte vegetation of (sub)atlantic, sandy, lowland regions during this century

SUMMARY.

• 1 Considerable changes in macrophyte vegetation can be noticed in 146 originally soft waters, when data on the recent aquatic vegetation are compared with historical information from the period 1900–60. Changes in nutrient status (N, P and C) and accumulation of organic material can be regarded as the operative factors.
• 2 The processes observed in soft waters are acidification, eutrophication and water hardening. Which process dominates depends on the type of soft water.
• 3 Acidification as well as eutrophication of water bodies may ultimately result in the total disappearance of all aquatic macrophytes, with the exception of the floating-leaved nymphaeids Nymphaea alba L. and Nuphar lutea (L.) Sm. Observed successional stages are described and summarized.