Denitrification is a critical process that can alleviate the effects of excessive nitrogen availability in aquatic ecosystems subject to eutrophication. An important part of denitrification occurs in benthic systems where bioturbation by meiofauna (invertebrates <1 mm) and its effect on element cycling are still not well understood. Here we study the quantitative impact of meiofauna populations of different abundance and diversity, in the presence and absence of macrofauna, on nitrate reduction, carbon mineralization and methane fluxes. In sediments with abundant and diverse meiofauna, denitrification is double that in sediments with low meiofauna, suggesting that meiofauna bioturbation has a stimulating effect on nitrifying and denitrifying bacteria. However, high meiofauna densities in the presence of bivalves do not stimulate denitrification, while dissimilatory nitrate reduction to ammonium rate and methane efflux are significantly enhanced. We demonstrate that the ecological interactions between meio-, macrofauna and bacteria are important in regulating nitrogen cycling in soft-sediment ecosystems.

ABSTRACT(1) The succession of primary producer communities in coastal lagoons is analysed in the light of the regime shift theory. Pristine coastal lagoons are considered to be dominated by extensive meadows of seagrass species, which are assumed to take advantage of nutrient supply from sediments. An increasing nutrient input is thought to favour phytoplankton and/or epiphytic micro-, macroalgae as well as opportunistic ephemeral macroalgae that coexist with seagrasses. In the latest stages of this succession, the imbalance of phosphorus to nitrogen ratio can favour macroalgal, cyanobacteria and/or picoplankton blooms, often causing dystrophy.(2) The primary causes of shifts and succession in the macrophyte community are nutrient loadings, mainly nitrogen, as well as changes in coastal hydrology or interactions between them. To some extent, in very shallow choked lagoons, benthic vegetation is mainly controlled by loading rates, while in open deep estuaries hydromorphological factors predominate.(3) External stressors/perturbations cause an amplification in benthic biogeochemical processes, e.g. wide variations in primary productivity and dark respiration, with large oscillations in oxygen and sulphide concentrations. Altered biogeochemical processes can determine positive feedbacks inducing a shift from pristine to altered macrophyte communities, which in turn amplify the perturbation until the shift becomes irreversible.(4) Macrophyte typology, organic matter composition and sedimentary geochemistry are primary factors in controlling feedbacks and shifts. For example, the sedimentary buffering capacity of iron controls sulphide and phosphates, while nitrogen cycling is mainly controlled by primary producers -microbial process interactions.(5) The alternative states which occur through the transition from pristine to modified primary producer communities can also be viewed as a sequence of stable states with different degrees of embedded information and with different ecological functions.

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Nutrient reduction and the improvement of bottom water oxygen concentrations are thought to be key factors in the recovery of eutrophic aquatic ecosystems. The effects of reoxygenation and bioturbation of natural hypoxic sediments in the Baltic Sea were studied using a mesocosm experiment. Anoxic sediment box cores were collected from 100 m depth in Kanholmsfjärden (Stockholm Archipelago) and maintained in flow-through mesocosms with 3 treatments: (1) hypoxic: supplied with hypoxic water; (2) 3− and H 4 SiO 4 , and rates of nitrate ammonification (DNRA), denitrification and anammox were determined. Phosphate was taken up by the sediment in all treatments, and the uptake was highest in the normoxic treatment with Marenzelleria. Normoxic conditions stimulated the denitrification rate by a factor of 5. Denitrification efficiency was highest under normoxia (50%), intermediate in bioturbated sediments (16%), and very low in hypoxic sediments (4%). The shift from hypoxic to normoxic conditions resulted in a significantly higher retention of NH 4 + , H 4 SiO 4 and Mn 2+ in the sediment, but the bioturbation by Marenzelleria reversed this effect. Results from our study suggest that bioturbation by Marenzelleria stimulates the exchange of solutes between sediment and bottom water through irrigation and enhances bacterial sulfate reduction in the burrow walls. The latter may have a toxic effect on nitrifying bacteria, which, in turn, suppresses denitrification rates.

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Wetland ecosystems in agricultural areas often become progressively more isolated from main water bodies. Stagnation favors the accumulation of organic matter as the supply of electron acceptors with water renewal is limited. In this context it is expected that nitrogen recycling prevails over nitrogen dissipation. To test this hypothesis, denitrification rates, fluxes of dissolved oxygen (SOD), inorganic carbon (DIC) and nitrogen and sediment features were measured in winter and summer 2007 on 22 shallow riverine wetlands in the Po River Plain (Northern Italy). Fluxes were determined from incubations of intact cores by measurement of concentration changes or isotope pairing in the case of denitrification. Sampled sites were eutrophic to hypertrophic; 10 were connected and 12 were isolated from the adjacent rivers, resulting in large differences in nitrate concentrations in the water column (from \5 to 1,133 lM). Benthic metabolism and denitrification rates were investigated by two overarching factors: season and hydrological connectivity. SOD and DIC fluxes resulted in respiratory quotients greater than one at most sampling sites. Sediment respiration was coupled to both ammonium efflux, which increased from winter to summer, and nitrate consumption, with higher rates in river-connected wetlands. Denitrification rates measured in river-connected wetlands (35-1,888 lmol N m -2 h -1 ) were up to two orders of magnitude higher than rates measured in isolated wetlands (2-231 lmol N m -2 h -1 ), suggesting a strong regulation of the process by nitrate availability. These rates were also significantly higher in summer (9-1,888 lmol N m -2 h -1 ) than in winter (2-365 lmol N m -2 h -1 ). Denitrification supported by water column nitrate (D W ) accounted for 60-100% of total denitrification (Dtot); denitrification coupled to nitrification (D N ) was probably controlled by limited oxygen availability within sediments. Denitrification efficiency, calculated as the ratio between N removal via denitrification and N regeneration, and the relative role of denitrification for organic matter oxidation, were high in connected wetlands but not in isolated sites. This study confirms the importance of restoring hydraulic connectivity of riverine wetlands for the maintenance of important biogeochemical functions such as nitrogen removal via denitrification.

Organic enrichment may deeply affect benthic nitrogen (N) cycling in macrophyte meadows, either promoting N loss or its recycling. This depends upon the plasticity of plants and of the associated microbial communities, as those surrounding the rhizosphere. Rates of denitrification, dissolved inorganic N fluxes and N uptake were measured in sediments vegetated by the submerged macrophyte Vallisneria spiralis L. under increasing organic matter loads. The aim was to investigate how the combined N assimilation and denitrification, which subtract N via temporary retention and permanent removal, respectively, do vary along the gradient. Results showed that V. spiralis meadows act as regulators of benthic N cycling even in organic enriched sediments, with negative feedbacks for eutrophication. A moderate organic load stimulates N uptake and denitrification coupled to nitrification in the rhizosphere. This is due to a combination of weakened competition between macrophytes and N cycling bacteria and enhanced radial oxygen loss by roots. An elevated organic enrichment affects N uptake due to hostile conditions in pore water and plant stress and impairs N mineralisation and its removal via denitrification coupled to nitrification. However, the loss of plant performance is almost completely compensated by increased denitrification ofwater column nitrate, resulting in a shift between the relative relevance of temporary and permanent N removal processes

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