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 In 41 of them, Littorella had been replaced by submerged Juncus bulbosus L. and/or Sphagnum spp.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.     

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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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.     

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.     

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.     

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.     

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.     

Higher nitrate-reducer diversity in macrophyte-colonized compared to unvegetated freshwater sediment

Freshwater macrophytes stimulate rhizosphere-associated coupled nitrification–denitrification and are therefore likely to influence the community composition and abundance of rhizosphere-associated denitrifiers and nitrate reducers. Using the narG gene, which encodes the catalytic subunit of the membrane-bound nitrate reductase, as a molecular marker, the community composition and relative abundance of nitrate-reducing bacteria were compared in the rhizosphere of the freshwater macrophyte species Littorella uniflora and Myriophyllum alterniflorum to nitrate-reducing communities in unvegetated sediment. Microsensor analysis indicated a higher availability of oxygen in the rhizosphere compared to unvegetated sediment, with a stronger release of oxygen from the roots of L. uniflora compared to M. alterniflorum. Comparison of narG clone libraries between samples revealed a higher diversity of narG phylotypes in association with the macrophyte rhizospheres compared to unvegetated sediment. Quantitative PCR targeting narG- and 16S rRNA-encoding genes pointed to a selective enrichment of narG gene copies in the rhizosphere. The results suggested that the microenvironment of macrophyte rhizospheres, characterized by the release of oxygen and labile organic carbon from the root system, had a stimulating effect on the diversity and relative abundance of rhizosphere-associated nitrate reducers.     

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.     

Leaf production and nitrogen and phosphorus tissue content of Littorella uniflora (L.) Aschers. In relation to nitrogen and phosphorus enrichment of the sediment in oligotrophic Lake Hampen,Denmark

Littoral sediments with a dense population of Littorella uniflora (L.) Ascers. were artificially enriched with nitrogen and phosphorus. Responses to increasing levels of nutrients were recorded as leaf production and tissue nutrient content. Phosphorus enrichment resulted in increasing P concentrations in all plant fractions (leaves, stem and roots), whereas only leaves and roots showed increasing N concentrations when inorganic N was added. Leaf production increased significantly after increasing the sediment P level 2, 3 and 5 times. Nitrogen did not affect leaf production. The tissue content of total N and P in control plants averaged 3.29 and 0.28%, respectively. Thus the critical P level in L. uniflora seems to be about 0.28%, which is higher than previously published values. It was also found that L. uniflora translocates phosphorus from old to younger leaves.     

High resistance of oligotrophic isoetid plants to oxic and anoxic dark exposure

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Ecological effects of lime treatment of acidified lakes and rivers in Sweden

Effects on the aquatic biota of lime (CaCO₃) application in acidified lakes and streams were studied in a number of waters. After treatment, lime-sensitive species of mosses (Sphagnum spp.) decreased, but species such as Potamogeton natans and Myriophyllum alterniflorum seemed to be favoured. A few years after liming species composition and diversity of phytoplankton, zooplankton and benthic insect larvae were almost identical to that found in oligotrophic and non-acid lakes. Molluscs and benthic crustaceans may have difficulties recolonizing. Reproduction of remaining species of fish was successful as soon as pH increased. High survival of larvae and fry can result in some extremely rich year classes with slow individual growth. In most cases restocking of depleted fish stocks was successful.     

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

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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.     

Benthic response to ammonium pulses in a tropical lagoon: implications for coastal environmental processes

In New Caledonia, the benthic communities living in the coral reef lagoon around Noumea city are subjected to regular shifts from oligotrophic conditions typical of lagoon waters to nutrient enrichment due to waste water inputs. The influence of ammonium pulses on microphytobenthos production was experimentally tested under varying light intensities in the vicinity of Noumea. Benthic oxygen, ammonium and silicon fluxes at the sediment-water interface were measured in situ using benthic enclosures. Three ammonium concentrations were tested. Gross primary production was doubled with a 13.8 μmol l⁻¹ ammonium concentration increase. Fitted PI curves showed that maximum production (F_max) was linearly related to ammonium concentration, but not the optimal irradiance (I_k). Silicon fluxes were characterized by dissolution in the absence of light, a process that declined with increasing illumination. These results were attributed to microphytobenthos activity, mainly diatoms that are nutrient-limited and strongly reactive to ammonium inputs. Production may result from a multiplication of cells, but migration up to the water sediment interface may also be involved. Oxygen consumption was also significantly influenced by ammonium concentration as a positive linear relationship with added ammonium concentration was established. Even during short-term experiments, ammonium enrichment stimulated photoautotrophic production, increasing the energy available to heterotrophs. Furthermore, microbenthic activities as well as nitrate production were increased by ammonia-oxidizing bacteria able to grow chemolithotrophically at the expense of oxygen. Therefore, in the study area, pulses of urban waste waters resulted in a decrease of plant-related autotrophy in benthic communities.     

Heterotrophic utilization and decomposition of extracellular carbon released by the aquatic angiosperm Littorella uniflora (L.) aschers

The heterotrophic assimilation of extracellular organic carbon (EOC) released by Littorella uniflora (L.) Aschers. was assayed. Utilization and decompostion of EOC by heterotrophs in the periphyton community were measured in a two-chamber open flow-system and, by the bacterioplankton, in bottle incubations. Under the experimental conditions 12–30% of the released EOC was metabolized in the periphyton community. This corresponded to a theoretical removal rate of 72–100% h^?1. The bacterioplankton assimilated EOC more slowly (1% h^?1). These results show that the EOC (mainly small molecules < 700 Daltons) was highly labile and accessible to heterotrophic microorganisms. The EOC released by submerged macrophytes in Lake Kalgaard is distributed approximately 20% to the periphyton community, 20% to the bacterioplankton, and 60% to the sediment.     

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.     

Populations of Myriophyllum alterniflorum L. as bioindicators of pollution in acidic to neutral rivers in the Limousin region

Myriophyllum alterniflorum D.C. (Haloragaceae) was studied in the acidic to neutral rivers Vienne and Gartempe (Limousin region, France). Two complementary levels of organisation were evaluated to determine their usefulness for pollution assessment: morphological traits of river populations, and eco-physiological responses to ammonium enrichment in indoor experiments. Inter-node length, stem diameter and leaf length were measured and their ratio calculated for 100 selected plants collected in 20 homogeneous areas. Using multidimensional analysis, their ecological significance for trophic assessment was demonstrated: stem diameter was linked to conductivity, and leaf length/stem diameter ratio to phosphates. Nitrate reductase activity was measured in three populations (upstream, median and downstream populations) after experimental enrichment with ammonium. Ammonium enrichment decreased activity starting at very low concentrations. However, inhibition levels depended on tested populations, with upstream populations being less sensitive due to natural ammonium content in water. The differences could be explained by an eco-adaptation to the trophic level of water. The relationships between the two levels of organisation are discussed. These results could help to distinguish different ecotypes using Myriophyllum alterniflorum.     

CAM-like Photosynthesis in Littorella uniflora (L.) Aschers.: The Role of Humidity

The shoreweed Littorella uniflora (L.) Aschers. possesses twogrowth forms differing from each other both in morphologicaland ecological features. In two recent publications, oppositeresults have been reported on the occurrence of a CAM-like carbonpathway in the terrestrial form of L. uniflora. Experimentalanalysis in this paper reveals that the induction of diel acidmetabolism requires saturated air humidity, conditions whichrarely occur in nature. Littorella uniflora (L.) Aschers., CAM-like photosynthesis, malic acid, relative humidity     

Degraded Softwater Lakes: Possibilities for Restoration

In the Netherlands, the characteristic flora of shallow softwater lakes has declined rapidly as a consequence of eutrophication, alkalization and acidification. The sediment of most lakes has become nutrient rich and anaerobic. We expected that, if a vital seed bank was still present, restoration of the original water quality and sediment conditions would lead to the return of softwater macrophytes. The restoration of 15 degraded, shallow, softwater lakes in the Netherlands was monitored from 1983 to 1998. In eutrophied as well as in acidified lakes, removal of accumulated organic matter from the sediment and shores was followed by rapid recolonization of softwater macrophytes present in the seedbank. After isolation from alkaline water and subsequent mud removal, this recovery was also observed in alkalized lakes. Further development of softwater vegetation correlated strongly with the water quality. When renewed eutrophication was successfully prevented, softwater macrophytes could expand. However, in acidified lakes, Juncus bulbosus and Sphagnum species became dominant after restoration. Liming of an acidified lake was followed by re‐acidification within 3 years. Recolonization by softwater macrophytes was inhibited by high turbidity of the water column and spreading of large helophytes on the shore. As an alternative, controlled inlet of alkaline, nutrient‐poor groundwater was studied in a few lakes. The pH of those lakes increased, the carbon and nitrogen availability decreased and softwater macrophytes returned. Successful restoration has contributed considerably to maintaining biodiversity in softwater lakes in the Netherlands.