Food webs in isolation: The food-web structure of a freshwater reservoir with armoured shores in a former coastal bay area.

Many shallow coastal bays have been closed off from the sea to mitigate the risk of flooding, resulting in coastal reservoir lakes with artificial armoured shorelines. Often these enclosed ecosystems show a persistent decline in biodiversity and ecosystem services, which is likely reflected in their food-web structure. We therefore hypothesize that the food webs of coastal reservoir lakes with armoured shorelines (1) consist of relatively few species with a low food-web connectance and short food chains, and (2) are mainly fuelled by autochthonous organic matter produced in the pelagic zone. To investigate these two hypotheses, we used stable-isotope analysis to determine the food-web structure of lake Markermeer (The Netherlands), a large reservoir lake with armoured shorelines in a former coastal bay area. Contrary to expectation, connectance of the food web in lake Markermeer was comparable to other lakes, while food-chain length was in the higher range. However, the trophic links revealed that numerous macroinvertebrates and fish species in this constructed lake exhibited omnivorous feeding behaviour. Furthermore, in line with our second hypothesis, primary consumers heavily relied on pelagically derived organic matter, while benthic primary production exerted only a minor and seasonal influence on higher trophic levels. Stable-isotope values and the C:N ratio of sediment organic matter in the lake also aligned more closely with phytoplankton than with benthic primary producers. Moreover, terrestrial subsidies of organic matter were virtually absent in lake Markermeer. These findings support the notion that isolation of the lake through shore armouring and the lack of littoral habitats in combination with persistent resuspension of sediments have affected the food web. We argue that restoration initiatives should prioritize the establishment of land-water transition zones, thereby enhancing habitat diversity, benthic primary production, and the inflow of external organic matter while preserving pelagic primary production.

Over forty years of lowland stream restoration: Lessons learned?

Stream restoration efforts have increased, but the success rate is still rather low. The underlying reasons for these unsuccessful restoration efforts remain inconclusive and need urgent clarification. Therefore, the aim of the present study was to evaluate over 40 years of stream restoration to fuel future perspectives. To this purpose we evaluated the influence of policy goals on stream restoration efforts, biophysical restoration objectives, restoration measures applied including the scale of application and monitoring efforts. Information was obtained from five stream restoration surveys that were held among the regional water authorities in the Netherlands over the last 40 years and from an analysis of the international scientific publications on stream restoration spanning the same time period. Our study showed that there was a considerable increase in stream restoration efforts, especially motivated by environmental legislation. However, proper monitoring of the effectiveness of the measures was often lacking. Furthermore, a mismatch between restoration goals and restoration measures was observed. Measures are still mainly focused on hydromorphological techniques, while biological goals remain underexposed and therefore need to be better targeted. Moreover, restoration practices occur mainly on small scales, despite the widely recognized relevance of tackling multiple stressors acting over large scales for stream ecosystem recovery. In order to increase the success rate of restoration projects, it is recommended to improve the design of the accompanying monitoring programmes, allowing to evaluate, over longer time periods, if the measures taken led to the desired results. Secondly, we advise to diagnose the dominant stressors and plan restoration measures at the appropriate scale of these stressors, generally the catchment scale.

Responses of macroinvertebrate communities to land use specific sediment food and habitat characteristics in lowland streams.

The input of land use specific organic matter into lowland streams may impact sediment characteristics in terms of food resources and habitat structure, resulting in differences in macroinvertebrate community composition. Therefore, we investigated to what extent land use specific sediment food and habitat characteristics structure macroinvertebrate communities. To this purpose linear multiple regression models were constructed, in which macroinvertebrate biotic indices were considered as response variables and sediment characteristics as predictor variables, analysed in 20 stream stretches running through five different land use types. Sediment characteristics and macroinvertebrate community composition were land use specific. The carbon/nitrogen (C/N) ratio, woody debris substrate cover and the origin of fatty acids influenced macroinvertebrate community composition. Shannon-Wiener diversity was better explained by fatty acids origin, such as in grassland streams, where a higher relative content of plant derived fatty acids related to a higher macroinvertebrate diversity. In cropland and wastewater treatment plant (WWTP) streams with a low C/N ratio and dominated by microbial derived fatty acids, higher abundances of Oligochaeta and Chironomus sp. were observed. Ephemeroptera, Plecoptera, and Trichoptera (EPT) richness was positively related to woody debris substrate cover, which only occurred in forest streams. Hence, macroinvertebrate community composition was influenced by the origin of the organic material, being either allochthonous or autochthonous and when autochthonous being either autotrophic or heterotrophic. It is therefore concluded that sediment food and habitat characteristics are key ecological filters.

Land use affects lowland stream ecosystems through dissolved oxygen regimes.

The aim of the present study was to assess the impact of surrounding land use on the structure and functioning of lowland stream ecosystems. To this end, five different land use types were selected (forest, extensive grassland, intensive grassland, cropland and wastewater treatment plant) each represented by four replicate streams, in which diel dissolved oxygen concentrations were recorded, sediment and water quality parameters were measured and macroinvertebrate community composition was determined. Chironomus sp., Oligochaeta and Gastropoda dominated the cropland and wastewater treatment plant (WWTP) streams, while Plecoptera and most Trichoptera only occurred in forest and extensive grassland streams. Forest streams communities were related to a high oxygen saturation, a high C/N ratio in the sediment and woody debris and coarse particulate organic matter (CPOM) substrate cover. Macroinvertebrate communities in cropland and WWTP streams were related to a low oxygen saturation in water and sediment and high concentrations of dissolved nitrogen, phosphorus and carbon. It is concluded that land use specific impacts on lowland streams are likely exerted via fine sediment accumulation in deposition zones, affecting oxygen regimes, sediment oxygen demand and stream metabolism, ultimately changing macroinvertebrate community composition. This study supports therefore the importance of including the catchment scale in ecological stream quality assessments, combining structural and functional endpoints.

Atmospheric Electricity Influencing Biogeochemical Processes in Soils and Sediments.

The Earth's subsurface represents a complex electrochemical environment that contains many electro-active chemical compounds that are relevant for a wide array of biologically driven ecosystem processes. Concentrations of many of these electro-active compounds within Earth's subsurface environments fluctuate during the day and over seasons. This has been observed for surface waters, sediments and continental soils. This variability can affect particularly small, relatively immobile organisms living in these environments. While various drivers have been identified, a comprehensive understanding of the causes and consequences of spatio-temporal variability in subsurface electrochemistry is still lacking. Here we propose that variations in atmospheric electricity (AE) can influence the electrochemical environments of soils, water bodies and their sediments, with implications that are likely relevant for a wide range of organisms and ecosystem processes. We tested this hypothesis in field and laboratory case studies. Based on measurements of subsurface redox conditions in soils and sediment, we found evidence for both local and global variation in AE with corresponding patterns in subsurface redox conditions. In the laboratory, bacterial respiratory responses, electron transport activity and H2S production were observed to be causally linked to changes in atmospheric cation concentrations. We argue that such patterns are part of an overlooked phenomenon. This recognition widens our conceptual understanding of chemical and biological processes in the Earth's subsurface and their interactions with the atmosphere and the physical environment.

Connectivity and seasonality cause rapid taxonomic and functional trait succession within an invertebrate community after stream restoration.

General colonization concepts consent that a slow process of microhabitat formation and subsequent niche realization occurs during early stages after new habitat is released. Subsequently, only few species are able to colonize new habitat in the early onset of succession, while species richness increases steadily over time. Although most colonization studies have been performed in terrestrial ecosystems, running water ecosystems are equally or even more prone to colonization after disturbance due to their dynamic nature. We question how invertebrate succession patterns reconcile with general colonization concepts. With this study we provide insight into the colonization process in newly created lowland stream trajectories and answer how within-stream bio- and functional diversity develops over time. Our results show a rapid influx of species, with a wide range of functional traits, during the first season after water flow commenced. During more than two years of regular monitoring, immigration rates were highest in autumn, marking the effects of seasonality on invertebrate dispersal. Biodiversity increased while abundance peaks of species alternated between seasons. Moreover, also days since start of the experiment explains a considerable part of the variability for taxa as well as traits. However, the relative trait composition remained similar throughout the entire monitoring period and only few specific traits had significantly higher proportions during specific seasons. This indicates that first phase colonization in freshwater streams can be a very rapid process that results in a high biodiversity and a large variety of species functional characteristics from the early onset of succession, contradicting general terrestrial colonization theory.

Sediment composition mediated land use effects on lowland streams ecosystems.

Despite the widely acknowledged connection between terrestrial and aquatic ecosystems, the contribution of runoff to the sediment composition in lowland stream deposition zones and the subsequent effects on benthic invertebrates remain poorly understood. The aim of this study was therefore to investigate the mechanisms by which runoff affects sediment composition and macroinvertebrates in deposition zones of lowland stream ecosystems. To this end, sediment from runoff and adjacent instream deposition zones from streams with different land use was chemically characterized and the biological effects were assessed at the species, community and ecosystem level. Runoff and deposition zone sediment composition as well as biological responses differed clearly between forest and agricultural streams. The stream deposition zone sediment C/N ratio reflected the respective runoff sediment composition. Deposition zones in the forest stream had a higher C/N ratio in comparison to the agricultural streams. Growth of Hyalella azteca and reproduction of Asellus aquaticus were higher on forest stream sediment, whereas chironomids and worms suffered less mortality on the agricultural sediments containing only natural food. The forest stream deposition zones showed higher values for indices indicative of biological integrity and had a lower sediment oxygen demand. We concluded that agricultural land use affects lowland stream ecosystem deposition zones at the species, community and ecosystem level via altered food quality (C/N ratio) and higher oxygen demand of the sediment.

Cell kinetics during regeneration in the sponge Halisarca caerulea: how local is the response to tissue damage?

Sponges have a remarkable capacity to rapidly regenerate in response to wound infliction. In addition, sponges rapidly renew their filter systems (choanocytes) to maintain a healthy population of cells. This study describes the cell kinetics of choanocytes in the encrusting reef sponge Halisarca caerulea during early regeneration (0-8 h) following experimental wound infliction. Subsequently, we investigated the spatial relationship between regeneration and cell proliferation over a six-day period directly adjacent to the wound, 1 cm, and 3 cm from the wound. Cell proliferation was determined by the incorporation of 5-bromo-2'-deoxyuridine (BrdU). We demonstrate that during early regeneration, the growth fraction of the choanocytes (i.e., the percentage of proliferative cells) adjacent to the wound is reduced (7.0 ± 2.5%) compared to steady-state, undamaged tissue (46.6 ± 2.6%), while the length of the cell cycle remained short (5.6 ± 3.4 h). The percentage of proliferative choanocytes increased over time in all areas and after six days of regeneration choanocyte proliferation rates were comparable to steady-state tissue. Tissue areas farther from the wound had higher rates of choanocyte proliferation than areas closer to the wound, indicating that more resources are demanded from tissue in the immediate vicinity of the wound. There was no difference in the number of proliferative mesohyl cells in regenerative sponges compared to steady-state sponges. Our data suggest that the production of collagen-rich wound tissue is a key process in tissue regeneration for H. caerulea, and helps to rapidly occupy the bare substratum exposed by the wound. Regeneration and choanocyte renewal are competing and negatively correlated life-history traits, both essential to the survival of sponges. The efficient allocation of limited resources to these life-history traits has enabled the ecological success and diversification of sponges.

Resource niche overlap promotes stability of bacterial community metabolism in experimental microcosms.

Decomposition of organic matter is an important ecosystem process governed in part by bacteria. The process of decomposition is expected to benefit from interspecific bacterial interactions such as resource partitioning and facilitation. However, the relative importance of resource niche breadth (metabolic diversity) and resource niche overlap (functional redundancy) on decomposition and the temporal stability of ecosystem processes received little scientific attention. Therefore, this study aims to evaluate the effect of an increase in bacterial community resemblance on both decomposition and the stability of bacterial metabolism in aquatic sediments. To this end, we performed laboratory microcosm experiments in which we examined the influence of bacterial consortia differing in number and composition of species on bacterial activity (Electron Transport System Activity, ETSA), dissolved organic carbon production and wavelet transformed measurements of redox potential (Eh). Single substrate affinities of the individual bacterial species were determined in order to calculate the metabolic diversity of the microbial community. Results presented here indicate that bacterial activity and organic matter decomposition increase with widening of the resource niche breadth, and that metabolic stability increases with increasing overlap in bacterial resource niches, hinting that resource niche overlap can promote the stability of bacterial community metabolism.

Cell turnover and detritus production in marine sponges from tropical and temperate benthic ecosystems.

This study describes in vivo cell turnover (the balance between cell proliferation and cell loss) in eight marine sponge species from tropical coral reef, mangrove and temperate Mediterranean reef ecosystems. Cell proliferation was determined through the incorporation of 5-bromo-2'-deoxyuridine (BrdU) and measuring the percentage of BrdU-positive cells after 6 h of continuous labeling (10 h for Chondrosia reniformis). Apoptosis was identified using an antibody against active caspase-3. Cell loss through shedding was studied quantitatively by collecting and weighing sponge-expelled detritus and qualitatively by light microscopy of sponge tissue and detritus. All species investigated displayed substantial cell proliferation, predominantly in the choanoderm, but also in the mesohyl. The majority of coral reef species (five) showed between 16.1±15.9% and 19.0±2.0% choanocyte proliferation (mean±SD) after 6 h and the Mediterranean species, C. reniformis, showed 16.6±3.2% after 10 h BrdU-labeling. Monanchora arbuscula showed lower choanocyte proliferation (8.1±3.7%), whereas the mangrove species Mycale microsigmatosa showed relatively higher levels of choanocyte proliferation (70.5±6.6%). Choanocyte proliferation in Haliclona vansoesti was variable (2.8-73.1%). Apoptosis was negligible and not the primary mechanism of cell loss involved in cell turnover. All species investigated produced significant amounts of detritus (2.5-18% detritus bodyweight(-1)·d(-1)) and cell shedding was observed in seven out of eight species. The amount of shed cells observed in histological sections may be related to differences in residence time of detritus within canals. Detritus production could not be directly linked to cell shedding due to the degraded nature of expelled cellular debris. We have demonstrated that under steady-state conditions, cell turnover through cell proliferation and cell shedding are common processes to maintain tissue homeostasis in a variety of sponge species from different ecosystems. Cell turnover is hypothesized to be the main underlying mechanism producing sponge-derived detritus, a major trophic resource transferred through sponges in benthic ecosystems, such as coral reefs.

Laboratory algal bioassays using PAM fluorometry: effects of test conditions on the determination of herbicide and field sample toxicity.

Pulse Amplitude Modulation (PAM) fluorometry, based on chlorophyll a fluorescence, is a frequently used technique in algal bioassays to assess toxicity of single compounds or complex field samples. Several test conditions can influence the test results, and because a standardized test protocol is currently lacking, linking the results of different studies is difficult. Therefore, the aim of the present study was to gain insight into the effects of test conditions of laboratory algal bioassays using PAM fluorometry on the outcome of toxicity tests. To this purpose, we described the results from several pilot studies on test development in which information is provided on the effects of the main test factors during the pretest phase, the test preparation, the exposure period, and the actual measurement. The experiments were focused on individual herbicides and complex field samples and included the effects of culturing conditions, cell density, solvent concentration, exposure time, and the presence of actinic light. Several of these test conditions were found to influence the outcome of the toxicity test, and the presented information provides important background information for the interpretation of toxicity results and describes which test conditions should be taken into account when using an algal bioassay with PAM fluorometry. Finally, the application of PAM fluorometry in algal toxicity testing is discussed.

Hazard and risk of herbicides for marine microalgae.

Due to their specific effect on photosynthesis, herbicides pose a potential threat to coastal and estuarine microalgae. However, comprehensive understanding of the hazard and risk of these contaminants is currently lacking. Therefore the aim of the present study was to investigate the toxic effects of four ubiquitous herbicides (atrazine, diuron, Irgarol(®)1051 and isoproturon) and herbicide mixtures on marine microalgae. Using a Pulse Amplitude Modulation (PAM) fluorometry based bioassay we demonstrated a clear species and herbicide specific toxicity and showed that the current environmental legislation does not protect algae sufficiently against diuron and isoproturon. Although a low actual risk of herbicides in the field was demonstrated, monitoring data revealed that concentrations occasionally reach potential effect levels. Hence it cannot be excluded that herbicides contribute to observed changes in phytoplankton species composition in coastal waters, but this is likely to occur only occasionally.

Dynamics of natural populations of the dertitivorous mudsnail Potamopyrgus antipodarum (Gray) (Hydrobiidae) in two interconnected Lakes differing in trophic state.

Here we investigate the allocation of resources between growth and reproduction by surveying the dynamics of natural populations of the aquatic detritivorous mudsnail Potamopyrgus antipodarum from two interconnected lakes that differ in trophic state. The size distributions and reproductive output of the snail populations was analyzed monthly by field surveys spanning 3 years, and in a controlled microcosm experiment to evaluate the reproductive potential under laboratory conditions. Snails in the meso-oligotrophic lake showed reduced growth and a smaller size compared to snails in the eutrophic lake. However, the numbers of eggs and nearly-neonates per adult snail did not differ significantly between the two populations. It is speculated that P. antipodarum populations living under meso-oligotrophic conditions may consistently invest more internal energy in reproduction at the expense of growth and that food quantity may be an important driver for macro-invertebrate resource allocation in detrital food webs.

Changes in the planktonic microbial community during residence in a surface flow constructed wetland used for tertiary wastewater treatment.

Suspended particles are a major constituent of municipal wastewater and generally contain high levels of bacteria, including human pathogens. Discharge of these particles of anthropogenic nature can have profound effects on receiving aquatic ecosystems and mitigation of these effects requires additional polishing of treated municipal wastewater. Previously it was shown that surface flow constructed wetlands are effective in improving water quality by reducing the numbers of fecal indicator organisms. However, fecal indicator organisms represent only a minor fraction of the total planktonic bacterial community and knowledge on the effects of these constructed wetlands on the composition and functioning of the entire planktonic bacterial community is limited. The aim of this descriptive study was therefore to identify changes in the planktonic bacterial community during residence of secondary treated municipal wastewater in a full-scale surface flow constructed wetland. To this purpose water samples were taken in which the bacterial community composition and functioning were analyzed using FISH, DGGE and BIOLOG. Surprisingly, the bacterial abundance at the inflow of the constructed wetland was relatively low compared with more natural surface waters. However, the inflowing bacterial community showed high metabolic activity and functional diversity. During residence in the surface flow constructed wetland the bacterial abundance doubled, but decreased in metabolic activity and functional diversity. Shifts in the community composition indicate that these changes are related to turn-over of the bacterial community. The planktonic bacterial community in the effluent of the constructed wetland closely resembled natural bacterial communities in urban and agricultural ditches. Based on these observations we conclude that constructed wetlands are capable to mitigate possible impacts of the particle load in treated wastewaters by transforming the anthropological bacterial community to a bacterial community resembling more "natural" surface waters.

Suspended particle and pathogen peak discharge buffering by a surface-flow constructed wetland.

Constructed wetlands (CWs) have been shown to improve the water quality of treated wastewater. The capacity of CWs to reduce nutrients, pathogens and organic matter and restore oxygen regime under normal operating conditions cannot be extrapolated to periods of incidental peak discharges. The buffering capacity of CWs during peak discharges is potentially a key factor for water quality in the receiving waters. Therefore, the aim of the present study was to investigate the behaviour of peak discharges of suspended particles, (associated) physiochemical parameters and pathogenic organisms from a wastewater treatment plant (WWTP) in a full scale constructed wetland (CW). By mixing clarified water and sludge rich water from the settlement tank of the WWTP, the suspended particle concentration was increased for 8 h from ± 3.5 to ± 230 mg L(-1), and discharged into a full scale horizontal surface flow constructed wetland. An increase of suspended particle concentration following the peak discharge concurred with increases in turbidity and oxygen demand, total nutrient load (nitrogen, phosphorus and carbon) and pathogens (Escherichia coli and Enterococci). Temperature, pH, conductivity and dissolved nutrient concentrations (nitrogen, phosphorus and carbon) were however unaffected by the initial peak discharge. After retention in the unvegetated ponds (the first CW compartment) the applied suspended particle peak with a total load of 86.2 kg was reduced by >99%. Similar peak buffering was observed for the turbidity, oxygen demand and settable volume. Simultaneously dissolved nutrient concentrations increased, indicating partial mineralization of the suspended particles during retention in the unvegetated ponds. The peak buffering of pathogens was lower (40-84%), indicating differences in removal processes between other suspended particles and pathogens. The results indicated that the suspended particles were probably mostly removed by sedimentation and mineralization, where pathogens were more likely buffered by biofilm retainment, mortality and predation, mainly in reed ditches. After passing through the total CW the residuals of the suspended particle peak discharge were temporal increased concentrations of inorganic carbon (IC), NH(4) and E. coli (respectively 11%, 17% and 160% higher than steady state concentrations). The observations support the positive role of CWs for effective buffering of wastewater discharge peaks.

Dynamics of metal availability and toxicity in historically polluted floodplain sediments.

Many floodplains contain high concentrations of sediment associated contaminants that might be subjected to large changes in terms of mobility, transformation and bioavailability. Therefore, this study describes 1) changes in the redox conditions and the mobility of metals in artificially uncovered polluted floodplain sediments, 2) metal uptake by organisms and 3) colonization, succession and functioning of benthic algae on these sediments. Flooding caused long term changes in redox potential (Eh) profiles. In top layers strong gradients in redox potential established quickly, while in deeper layers changes occurred more gradually. The availability of copper as measured by Diffusive Gradients in Thinfilms (DGT) showed a consistent relationship with fluctuations in Eh. However, this relationship was restricted to deeper layers in the sediment. Within 1 week high amounts of total copper were immobilized. Differences in total copper concentrations between polluted and clean sediments became only partially apparent when comparing the available copper fraction (DGT-Cu). Introduced Tubifex shows only marginal differences in levels of accumulated Cu. Colonization, growth and succession of algal communities on polluted sediments was not impaired, most likely due to low bioavailability. It is concluded that changing environmental conditions, such as flooding, can result in stable chemical conditions with low a availability of metals and hence in a diminution of actual ecological risks.

Automated and continuous redox potential measurements in soil.

Redox potential (Eh) describes the electrical state of a matrix. In soils, Eh is an important parameter controlling the persistence of many organic and inorganic compounds. A popular, but also criticized, manual measuring method makes use of a small tip of Pt placed on a copper wire that is placed in the soil; a reference electrode is placed in the same soil at a fixed distance. Fluctuations in redox potential values measured in the soil can be very large and depth-dependent. This will be overlooked when making single-point measurements. We developed the datalogger Hypnos 2.0 for continuous redox potential and temperature measurements at various depths in the soil and without disturbance of the site. Hypnos is field-deployable, relatively cheap, and runs on batteries. The datalogger can use a "sleep mode" between sampling events. In sleep mode, there is no constant voltage on the Pt wire or the reference electrode, but there is only a short pulse during sampling. We did not measure an effect of this short pulse on the measured redox potential. In sandy soils in mesocosms and in a salt marsh soil we measured changes in the Eh as large as from -400 to +100 mV within 4 d, and daily cycles of 200 mV. Both absolute redox potential values and their diurnal variations were depth-dependent. Because single redox measurements are insufficient in describing redox conditions in some soil systems, Hypnos can be a powerful tool when studying the effects of fluctuating redox conditions on metal availability and pollutant degradation.

Combined effects of lowered oxygen and toxicants (copper and diazinon) on the mayfly Ephoron virgo.

In many large European rivers, the number of typical riverine insect species, such as mayflies, stoneflies, and caddisflies, is greatly reduced compared to historic records. This can no longer be explained by high concentrations of a relatively small number of dominant toxicants since many rivers have changed from heavily contaminated systems with a few selected key toxicants to systems with a complex contamination. This contamination consists of many substances in low concentrations coinciding with other unfavorable conditions, such as low oxygen concentrations. It was hypothesized that the joint adverse effects of such multiple stressors may be a steering factor in the distribution of riverine insect species. The aim of this study was therefore to determine the combined effects of toxicants and oxygen depletion. To this purpose, larvae of the indigenous riverine mayfly Ephoron virgo were exposed to two different model toxicants, copper and diazinon, under normoxia and hypoxia (50% air saturation) conditions. The median effective concentrations for mortality for copper were significantly lower in the hypoxia treatments than in the normoxia treatments. For diazinon, no differences were observed between two treatments differing in dissolved oxygen levels, and therefore we argue that interactions between multiple stressors may be compound specific. It is concluded that the combination of toxicants and lowered oxygen may have a stronger impact than can be expected based on the adverse effects of the individual factors and that standard toxicity tests may be insufficient alone to determine the impact of human activities on the ecological state of riverine communities. Instead, attention needs to be paid to more environmentally realistic nonoptimal conditions in toxicity testing to adequately fulfill the needs of ecological recovery programs.