In shallow‐water sediments, the combined action of microphytobenthos and bioturbating fauna may differentially affect benthic nutrient fluxes and exert a bottom‐up control of pelagic primary production. In many cases, the effects of microphytobenthos and macrofauna on nutrient cycling were studied separately, ignoring potential synergistic effects. We measured the combined effects of microphytobenthos and chironomid larvae on sediment–water fluxes of gas (O2, TCO2 and N2) and nutrients (NH4+, NO3−, NO2−, PO43− and SiO2) in shallow‐water sediments of a hypertrophic freshwater lagoon. Fluxes were measured in the light and in the dark in reconstructed sediments with low (L = 600 ind/m2), high (H = 1,800 ind/m2) and no (C) addition of chironomid larvae, after 3 weeks of pre‐incubation under light/dark regime to allow for microalgal growth. Besides flux measurements, pore water nutrient (NH4+, PO43− and SiO2) and dissolved metal concentrations (Fe2+ and Mn2+) were analysed and diffusive fluxes were calculated. Chironomid larvae increased sediment heterotrophy, by augmenting benthic O2 demand and TCO2 and N2 dark production. However, on a daily basis, treatments C and L were net O2 producing and N2 sinks while treatment H was net O2 consuming and N2 producing. All treatments were net C sink regardless of chironomid density. Microphytobenthos always affected benthic nutrient exchange, as significantly higher uptake or lower efflux was measured in the light compared with dark incubations. Theoretical inorganic N, P and Si demand by benthic microalgae largely exceeded both dark effluxes of NH4+, PO43− and SiO2 and their net uptake in the light, suggesting the relevance of N‐fixation, water column NO3− and solid‐phase associated P and Si as nutrient sources to benthic algae. Chironomid larvae had a minor effect on inorganic N and P fluxes while they significantly stimulated inorganic Si regeneration. Their bioturbation activity significantly altered pore water chemistry, with a major reduction in nutrient (highest for NH4+ and lowest for SiO2) and metal concentration. Underlying mechanisms are combinations of burrow ventilation and bioirrigation with stimulation of element‐specific processes as coupled nitrification–denitrification, co‐precipitation and inhibition of anaerobic paths such as Fe3+ or Mn4+ reduction or re‐oxidation of their end products. The combined activity of benthic algae and chironomid larvae may significantly attenuate internal nutrient recycling in shallow eutrophic ecosystems, and contribute to the control of pelagic primary production.
The Curonian Lagoon is Europe's largest lagoon and one of the most seriously impacted by harmful blooms of cyanobacteria. Intensive studies over the past 20 years have allowed us to identify the major drivers determining the composition and spatial extent of hyperblooms in this system. We summarize and discuss the main outcomes of these studies and provide an updated, conceptual scheme of the multiple interactions between climatic and hydrologic factors, and their influence on internal and external processes that promote cyanobacterial blooms. Retrospective analysis of remote sensed images demonstrated the variability of blooms in terms of timing, extension and intensity, suggesting that they occur only under specific circumstances. Monthly analysis of nutrient loads and stoichiometry from the principal tributary (Nemunas River) revealed large interannual differences in the delivery of key elements, but summer months were always characterized by a strong dissolved inorganic N (and Si) limitation, that depresses diatoms and favors the dominance of cyanobacteria. Cyanobacteria blooms occurred during high water temperatures, long water residence time and low-wind conditions. The blooms induce transient (night-time) hypoxia, which stimulates the release of iron-bound P, producing a positive feedback for blooms of N-fixing cyanobacteria. Consumermediated nutrient recycling by dreissenid mussels, chironomid larvae, cyprinids and large bird colonies, may also affect P availability, but their role as drivers of cyanobacteria blooms is understudied.
Bioturbation studies have generally analyzed small and abundant organisms while the contribution to the benthic metabolism by rare, large macrofauna has received little attention. We hypothesize that large, sporadic bivalves may represent a hot spot for benthic processes due to a combination of direct and indirect effects as their metabolic and bioturbation activities. Intact riverine sediments with and without individuals of the bivalve Sinanodonta woodiana were collected in a reach with transparent water, where the occurrence of the mollusk was clearly visible. The bivalve metabolism and its effects on sedimentary fluxes of dissolved gas and nutrients were measured via laboratory incubations of intact cores under controlled conditions. S. woodiana contributed significantly to O2 and TCO2 benthic fluxes through its respiration and to (Formula presented.), SRP and SiO2 regeneration via its excretion. The bivalve significantly stimulated also microbial denitrification and determined a large efflux of CH4, likely due a combination of bioturbation and biodeposition activities or to anaerobic metabolism within the mollusk gut. This study demonstrates that a few, large individuals of this bivalve produce significant effects on aerobic and anaerobic benthic metabolism and nutrient mobilization. Random sediment sampling in turbid waters seldom catches these important effects due to low densities of large fauna
The regulation of benthic nitrogen (N) cycling by multiple interactions among bacteria, macrofauna, and primary producers is poorly understood. We hypothesized that a biodiverse benthic system should better exploit the benthic N‐availability and retain N than a simpler one. Retention occurs by avoiding losses both to the water column via increased recycling and to the atmosphere via decreased N2 fluxes and by limiting energy‐costly processes as N‐fixation. We also hypothesized that primary producer‐bacterial competition is reduced in the presence of macrofauna due to mobilization of refractory N pools. To this purpose, the effects of two bioturbators (the detritivorous Sparganophilus tamesis and the filter‐feeding Corbicula spp.) and two primary producer growth forms (the rooted macrophyte Vallisneria spiralis and microphytobenthos) on benthic N cycling were studied. An array of N‐processes were measured along a complexity gradient (from bare sediments to all combinations of the above mentioned organisms), and experimental outcomes were analyzed via ecological network analysis (ENA). This suite of algorithms, applied to the microscale, revealed differential partitioning of N fluxes among bare sediments (highest denitrification rates), sediments with macrofauna (highest recycling), and sediments with rooted plants (highest N‐fixation). N2 losses and inputs were significantly reduced when all components were represented, and N requirements by primary producers were to a large extent supported by the activity of macrofauna. Ecological interactions in biodiverse benthic systems promoted an efficient exploitation of sedimentary N pools, increased the coupling between recycling and uptake, and maximized N use efficiency at the expenses of losses and imports.
Aquatic macrophytes modify the sediment biogeochemistry via radial oxygen loss (ROL) from their roots. However, the variation in ROL and its implication for nutrient availability remains poorly explored. Here, we use planar O2 optodes to investigate the spatial heterogeneity of oxic niches within the rhizosphere of Vallisneria spiralis and their alteration following variable light and ambient O2 levels. The effect of ROL on NH4+ and PO43− distribution in the rhizosphere was evaluated by a combination of 15N isotopic techniques, 2D sampling, and electron microscopy. A single specimen of V. spiralis could maintain an oxidised sediment volume of 41–47 cm3 and 10–27 cm3 in the rhizosphere at 100% and 38% dissolved oxygen saturation in the overlying water, respectively. Whatever the environmental conditions, the ROL was, however, very heterogeneous and dependent on root age and architecture of the root system. ROL stimulated the coupling between denitrification and nitrification in the sediment both under dark (+25 μmol N‐N2 m−2 hr−1) and light (+70 μmol N‐N2 m−2 hr−1) conditions. This, in combination with plant uptake, contributed to intense removal of NH4+ from the pore water. Similarly, PO43− was highly depleted in the rhizosphere. The detection of Fe‐P plaques on the roots surface indicated substantial entrapment of P as a consequence of ROL. The extensive spatio‐temporal heterogeneity of oxic and anoxic conditions ensured that aerobic and anaerobic processes co‐occurred in the rhizosphere and this presumably reduced potential nutrient limitation while maximising plant fitness in an otherwise hostile reduced environment.
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