Seasonal oxygen,nitrogen and phosphorus benthic cycling along an impacted Baltic Sea estuary: regulation and spatial patterns

The regulatory roles of temperature, eutrophication and oxygen availability on benthic nitrogen (N) cycling and the stoichiometry of regenerated nitrogen and phosphorus (P) were explored along a Baltic Sea estuary affected by treated sewage discharge. Rates of sediment denitrification, anammox, dissimilatory nitrate reduction to ammonium (DNRA), nutrient exchange, oxygen (O₂) uptake and penetration were measured seasonally. Sediments not affected by the nutrient plume released by the sewage treatment plant (STP) showed a strong seasonality in rates of O₂ uptake and coupled nitrification–denitrification, with anammox never accounting for more than 20 % of the total dinitrogen (N₂) production. N cycling in sediments close to the STP was highly dependent on oxygen availability, which masked temperature-related effects. These sediments switched from low N loss and high ammonium (NH₄ ⁺) efflux under hypoxic conditions in the fall, to a major N loss system in the winter when the sediment surface was oxidized. In the fall DNRA outcompeted denitrification as the main nitrate (NO₃ ^?) reduction pathway, resulting in N recycling and potential spreading of eutrophication. A comparison with historical records of nutrient discharge and denitrification indicated that the total N loss in the estuary has been tightly coupled to the total amount of nutrient discharge from the STP. Changes in dissolved inorganic nitrogen (DIN) released from the STP agreed well with variations in sedimentary N₂ removal. This indicates that denitrification and anammox efficiently counterbalance N loading in the estuary across the range of historical and present-day anthropogenic nutrient discharge. Overall low N/P ratios of the regenerated nutrient fluxes impose strong N limitation for the pelagic system and generate a high potential for nuisance cyanobacterial blooms.     

Influence of Potamogeton pectinatus and microphytobenthos on benthic metabolism,nutrient fluxes and denitrification in a freshwater littoral sediment in an agricultural landscape: N assimilation versus N removal

The influence of Potamogeton pectinatus colonisation on benthic nitrogen dynamics was studied in the littoral zone of a lowland pit lake with high nitrate concentration (~200 μM). Our hypothesis was that in aquatic environments where nitrogen availability is not limiting, colonisation by rooted macrophytes changes the dynamics of the benthic nitrogen cycle, stimulating N assimilation and denitrification and increasing the system capacity to take up external nitrogen loads. To test this hypothesis, we quantified and compared seasonal variations of light and dark benthic metabolism, dissolved inorganic nitrogen (DIN) fluxes, denitrification and N assimilation rates in an area colonised by P. pectinatus and a reference site colonised by microphytobenthos. In both areas, the benthic system was net autotrophic and a sink for DIN (2,241–2,644 mmol m^?2 y^?1). Plant colonisation increased nitrogen losses via denitrification by 30% compared to the unvegetated area. In contrast to what is generally observed in coastal marine systems, where the presence of rooted macrophytes limits denitrification rates, under the very high nitrate concentrations in the studied lake, both denitrification (1,237–1,570 mmol m^?2 y^?1) and N assimilation (1,039–1,095 mmol m^?2 y^?1) played important and comparable roles in the removal of DIN from the water column.     

Controls of Benthic Nitrogen Fixation and Primary Production from Nutrient Enrichment of Oligotrophic, Arctic Lakes

We examined controls of benthic dinitrogen (N₂) fixation and primary production in oligotrophic lakes in Arctic Alaska, Toolik Field Station (Arctic Long-Term Ecological Research Site). Primary production in many oligotrophic lakes is limited by nitrogen (N), and benthic processes are important for whole-lake function. Oligotrophic lakes are increasingly susceptible to low-level, non-point source nutrient inputs, yet the effects on benthic processes are not well understood. This study examines the results from a whole-lake fertilization experiment in which N and P were added at a relatively low level (4 times natural loading) in Redfield ratio to a shallow (3 m) and a deep (20 m) oligotrophic lake. The two lakes showed similar responses to fertilization: benthic primary production and respiration (each 50–150 mg C m^?2 day^?1) remained the same, and benthic N₂ fixation declined by a factor of three- to fourfold by the second year of treatment (from ~0.35 to 0.1 mg N m^?2 day^?1). This showed that the response of benthic N₂ fixation was de-coupled from the nutrient limitation status of benthic primary producers and raised questions about the mechanisms, which were examined in separate laboratory experiments. Bioassay experiments in intact cores also showed no response of benthic primary production to added N and P, but contrasted with the whole-lake experiment in that N₂ fixation did not respond to added N, either alone or in conjunction with P. This inconsistency was likely a result of nitrogenase activity of existing N₂ fixers during the relative short duration (9 days) of the bioassay experiment. N₂ fixation showed a positive saturating response when light was increased in the laboratory, but was not statistically related to ambient light level in the field, leading us to conclude that light limitation of the benthos from increasing water-column production was not important. Thus, increased N availability in the sediments through direct uptake likely caused a reduction in N₂ fixation. These results show the capacity of the benthos in oligotrophic systems to buffer the whole-system response to nutrient addition by the apparent ability for significant nutrient uptake and the rapid decline in N₂ fixation in response to added nutrients. Reduced benthic N₂ fixation may be an early indicator of a eutrophication response of lakes which precedes the transition from benthic to water-column-dominated systems.     

The importance of benthic–pelagic coupling for marine ecosystem functioning in a changing world

Benthic–pelagic coupling is manifested as the exchange of energy, mass, or nutrients between benthic and pelagic habitats. It plays a prominent role in aquatic ecosystems, and it is crucial to functions from nutrient cycling to energy transfer in food webs. Coastal and estuarine ecosystem structure and function are strongly affected by anthropogenic pressures; however, there are large gaps in our understanding of the responses of inorganic nutrient and organic matter fluxes between benthic habitats and the water column. We illustrate the varied nature of physical and biological benthic–pelagic coupling processes and their potential sensitivity to three anthropogenic pressures – climate change, nutrient loading, and fishing – using the Baltic Sea as a case study and summarize current knowledge on the exchange of inorganic nutrients and organic material between habitats. Traditionally measured benthic–pelagic coupling processes (e.g., nutrient exchange and sedimentation of organic material) are to some extent quantifiable, but the magnitude and variability of biological processes are rarely assessed, preventing quantitative comparisons. Changing oxygen conditions will continue to have widespread effects on the processes that govern inorganic and organic matter exchange among habitats while climate change and nutrient load reductions may have large effects on organic matter sedimentation. Many biological processes (predation, bioturbation) are expected to be sensitive to anthropogenic drivers, but the outcomes for ecosystem function are largely unknown. We emphasize how improved empirical and experimental understanding of benthic–pelagic coupling processes and their variability are necessary to inform models that can quantify the feedbacks among processes and ecosystem responses to a changing world.     

Microbial carbon oxidation rates and pathways in sediments of two Tanzanian mangrove forests

Temporal and spatial variations in benthic metabolism and anaerobic carbon oxidation pathways were assessed in an anthropogenically impacted (Mtoni) and a pristine (Ras Dege) mangrove forest in Tanzania. The objectives were (1) to evaluate how benthic metabolism is affected by organic carbon availability; (2) to determine the validity of diffusive release of CO₂ as a measure benthic carbon oxidation; and (3) to assess the partitioning of anaerobic carbon pathways and factors controlling the availability of electron acceptors (e.g. oxidized iron). Microbial carbon oxidation measured as diffusive exchange of O₂ and CO₂ (32?C67 and 28?C115 mmol m^?2 day^?1, respectively) showed no specific temporal patterns. Low intertidal sediments at Mtoni fed by labile algal carbon of anthropogenic origin had higher diffusive CO₂ release than high intertidal sediments that primarily received less reactive mangrove detritus. Diffusive release of CO₂ apparently underestimated total sediment carbon oxidation due to CO₂ loss from deep sediments via emission through biogenic structures (i.e. crab burrows and pneumatophores) and porewater seepage into creeks. We propose that diffusive fluxes in the present mangrove sediments are roughly equivalent to depth-integrated reactions occurring in the upper 12 cm. Anaerobic carbon oxidation was dominated by FeR irrespective of anthropogenic influence in sediments where the oxidizing effects of biogenic structures increased the Fe(III) level. More than 80% of the anaerobic carbon oxidation in Mtoni and Ras Dege sediments was due to FeR when reactive Fe(III) exceeded 30 ??mol cm^?3. The anthropogenic influence at Mtoni was primarily noted as up to one order of magnitude higher denitrification than at Ras Dege, but this process always accounted for less than 1% of total carbon oxidation. It is noteworthy that organic and nutrient enrichment of anthropogenic origin in Mtoni has no measurable effect on microbial processes, other than carbon oxidation in the low intertidal area and denitrification throughout the forest, and indicates a strong resilience of mangrove environments towards disturbances.     

Nitrification and denitrification in a midwestern stream containing high nitrate: in situ assessment using tracers in dome-shaped incubation chambers

The extent to which in-stream processes alter or remove nutrient loads in agriculturally impacted streams is critically important to watershed function and the delivery of those loads to coastal waters. In this study, patch-scale rates of in-stream benthic processes were determined using large volume, open-bottom benthic incubation chambers in a nitrate-rich, first to third order stream draining an area dominated by tile-drained row-crop fields. The chambers were fitted with sampling/mixing ports, a volume compensation bladder, and porewater samplers. Incubations were conducted with added tracers (NaBr and either ¹⁵N[NO₃ ^?], ¹⁵N[NO₂ ^?], or ¹⁵N[NH₄ ⁺]) for 24–44 h intervals and reaction rates were determined from changes in concentrations and isotopic compositions of nitrate, nitrite, ammonium and nitrogen gas. Overall, nitrate loss rates (220–3,560 μmol N m^?2 h^?1) greatly exceeded corresponding denitrification rates (34–212 μmol N m^?2 h^?1) and both of these rates were correlated with nitrate concentrations (90–1,330 μM), which could be readily manipulated with addition experiments. Chamber estimates closely matched whole-stream rates of denitrification and nitrate loss using ¹⁵N. Chamber incubations with acetylene indicated that coupled nitrification/denitrification was not a major source of N₂ production at ambient nitrate concentrations (175 μM), but acetylene was not effective for assessing denitrification at higher nitrate concentrations (1,330 μM). Ammonium uptake rates greatly exceeded nitrification rates, which were relatively low even with added ammonium (3.5 μmol N m^?2 h^?1), though incubations with nitrite demonstrated that oxidation to nitrate exceeded reduction to nitrogen gas in the surface sediments by fivefold to tenfold. The chamber results confirmed earlier studies that denitrification was a substantial nitrate sink in this stream, but they also indicated that dissolved inorganic nitrogen (DIN) turnover rates greatly exceeded the rates of permanent nitrogen removal via denitrification.     

Net sediment N2 fluxes in a southern New England estuary: variations in space and time

Over the past three decades, Narragansett Bay has undergone various ecological changes, including significant decreases in water column chlorophyll a concentrations, benthic oxygen uptake, and benthic nutrient regeneration rates. To add to this portrait of change, we measured the net flux of N₂ across the sediment–water interface over an annual cycle using the N₂/Ar technique at seven sites in the bay for comparison with measurements made decades ago. Net denitrification rates ranged from about 10–90 μmol N₂–N m^?2 h^?1 over the year. Denitrification rates were not significantly different among sites and had no clear correlation with temperature. Net nitrogen fixation (?5 to ?650 μmol N₂–N m^?2 h^?1) was measured at three sites and only observed in summer (June–August). Neither denitrification nor nitrogen fixation exhibited a consistent relationship with sediment oxygen demand or with fluxes of nitrite, nitrate, ammonium, total dissolved inorganic nitrogen, or dissolved inorganic phosphate across all stations. In contrast to the mid-bay historical site where denitrification rates have declined, denitrification rates in the Providence River Estuary have not changed significantly over the past 30 years.     

Transport of microbial biomass through the North Inlet ecosystem

Tidal fluctuations and transports of total microbial biomass (measured as adenosine triphosphate [ATP]) were investigated at three marsh creeks comprising the major transfer points between the North Inlet marsh and the adjoining aquatic ecosystems. Two creeks, Town Creek and North Jones Creek, form the inlet mouth and are the only marsh-ocean exchange points. The third creek, South Jones Creek, connects to a brackish water embayment. The creeks were simultaneously sampled every 1.5 hours for 50 consecutive hours during neap tides (four tidal cycles) and 50 consecutive hours during spring tides of each season. At the inlet, ATP concentration fluctuated in phase with the tide during winter and fall and out of phase with the tide during the spring. Fluctuation patterns at South Jones Creek were irregular. The highest ATP concentrations were during the spring (mean=2.17 mg of ATP per m³) and the lowest concentrations were during the winter (mean=0.65 mg of ATP per m³). Net transports of ATP varied from tidal cycle to tidal cycle with regard to direction of transport (import or export) and magnitude. Net transports were small compared to large instantaneous transports and only 4 of 22 determinations of net transport were different from zero transport.     

Evaluation of microalgal consortia for treatment of primary treated sewage effluent and biomass production

The present investigation was aimed towards analyzing the potential of consortia of native filamentous microalgal strains (MC2), native unicellular microalgal strains (MC3), and selected microalgae from germplasm (MC1) in terms of nutrient removal, water quality improvement, and biomass production using primary treated sewage water. Highest NO₃-N (90 %) and PO₄-P (97.8 %) removal was obtained with MC2-inoculated sewage water. Highest decrease in total dissolved solids to 806 from 1,120 mg L^?1 and highest increase in dissolved oxygen of 9.0 from 0.4 mg L^?1 were obtained using MC2-inoculated sewage water on the sixth day. The biomass production was also highest in MC2 (1.07 g L^?1) followed by MC1 and MC3 (0.90 and 0.94 g L^?1, respectively) on the sixth day. The consortium of filamentous strains from native environment not only proved promising in nutrient removal efficiency but also led to enhanced biomass. The present study highlighted the utility of such a consortium for sewage wastewater treatment and the promise of sewage water as a growth medium for biomass production.     

MMHg production and export from intertidal sediments to the water column of a tidal lagoon (Arcachon Bay,France)

Hg cycling in biologically productive coastal areas is of special importance given the potential for bioaccumulation of monomethylmercury (MMHg) into aquatic organisms. Field experiments were performed during three different seasons in Arcachon Bay, a mesotidal lagoon (SW France), to assess the variability of the water column concentrations, sediment–water exchanges and potential formation and degradation of MMHg. The objectives were to evaluate the contribution of intertidal mudflats to MMHg production and the various pathways of Hg species export. Dissolved and bulk concentrations of Hg species in the water column downstream of tidal flats were measured throughout several tidal cycles. The Hg benthic fluxes at the sediment–water interface were determined by means of benthic chambers for three different stations. Hg methylation and demethylation potentials were determined in surficial sediments and the water column using isotopic tracers. The tidal surveys demonstrated that benthic remobilization of Hg occurs primarily in association with sediment erosion and advection during ebb tide. However, elevated dissolved Hg concentrations observed at low tide were found to be caused by a combination of pore-waters seeping, benthic fluxes and methylation in the water column. Benthic fluxes were more intense during late winter conditions (median MMHg and inorganic Hg (IHg) fluxes: 64 and 179 pmol m^?2 h^?1, respectively) and subsequently decreased in spring (median 0.7 and ?5 pmol m^?2 h^?1, respectively) and fall (median ?0.4 and ?1.3 pmol m^?2 h^?1, respectively). The trends in methylation and demethylation potentials were at the opposite of the fluxes, two times lower during winter than for spring or fall conditions. In this tidal environment, MMHg production in surface sediments and its subsequent release is estimated to be the major source of MMHg to the water column during winter and spring time. However, during the more productive summer period, the Hg methylation extent in the water column may be very significant and equivalent to the sediment contribution.     

Close and delayed benthic–pelagic coupling in coastal ecosystems: the role of physical constraints

We aimed to analyse the temporal scales of the variability of benthic ammonium flux using data from an estuarine bay (Alfacs Bay, N. W. Mediterranean). Several nitrogen (N) pools in the sediment, their reactivities and their associated fluxes were estimated. Decomposition of labile buried N (4.5 mol N m^-2) was found to cause an ammonium flux of 0.1 mmol N m^-2 d ^-1, referred to as background flux. The fluxes measured from bell-jar incubations were usually higher, between 2 and 6 mmol N m^-2 d ^-1, which reflects mineralization of recent sedimentation. A typical sedimentation event was found to account for 25 – 75 mmol m^-2 of freshly settled N, which should bring on an ammonium flux of about 1.7–5.0 mmol N m^-2 d ^-1, referred to as fast flux. The concordance between measured and computed benthic fluxes is associated with the coupling of benthic fluxes to production and sedimentation. Close benthic–pelagic coupling was observed in winter and early spring, while a delayed flux response to sedimentation, with transient variations of pore water ammonium profiles, showing surface peaks and decreasing concentrations with depth, was found in autumn. Structures, such as viscous layers, which develop over the sediment–water interface, were found to be essential to the regulation of benthic processes and to lead to transient variations of pore water nutrient concentrations and associated fluxes. The temporal scales of the benthic flux response to sedimentation were discussed in terms of the physical structures involved in decomposition (the bulk sediment, the viscous layers or the fresh settled layer), the processes controlling kinetics and diffusion laws. Several scenarios for the benthic- pelagic coupling in Alfacs Bay, in which local (estuarine circulation) and climatic components combine to yield the variability observed, were examined.     

Long-term nutrient enrichment elicits a weak density response by saltmarsh meiofauna

The effect of chronic nutrient enrichment on benthic meiofauna was examined in a whole-ecosystem experiment conducted in salt marshes in the Plum Island watershed of northeastern Massachusetts, USA. We compared abundances of total meiofauna, nematodes, copepods, ostracods, and annelids including Manayunkia aestuarina, in fertilized (where N and P was increased 15× in incoming tidal water throughout each growing season for 6 years) and reference marsh creeks to test for bottom-up responses. Although some responses to nutrient enrichment were evident, results did not match our expectations of strong increases in abundance. Variation in abundance between nutrient-enriched and reference creeks was detected in all three subhabitats but responses were inconsistent and variable over time, suggesting that natural variability was greater than variation induced by fertilization. Our results showed an overall weak negative correlation between meiofauna abundance and benthic microalgae (BMA) biomass partly because the BMA response to nutrient enrichment was relatively small and perhaps limited by grazing macrofauna and nekton. Our results suggest a better mechanistic understanding of the relationship between BMA and meiofaunal abundance is needed to fully understand how nutrient enrichment affects meiofauna.     

Interaction of benthic microalgae and macrofauna in the control of benthic metabolism, nutrient fluxes and denitrification in a shallow sub-tropical coastal embayment (western Moreton Bay, Australia)

Benthic biogeochemistry and macrofauna were investigated six times over 1 year in a shallow sub-tropical embayment. Benthic fluxes of oxygen (annual mean ?918 μmol O₂ m^?2 h^?1), ammonium (NH₄ ⁺), nitrate (NO₃ ^?), dissolved organic nitrogen, dinitrogen gas (N₂), and dissolved inorganic phosphorus were positively related to OM supply (N mineralisation) and inversely related to benthic light (N assimilation). Ammonium (NH₄ ⁺), NO₃ ^? and N₂ fluxes (annual means +14.6, +15.9 and 44.6 μmol N m^?2 h^?1) accounted for 14, 16 and 53 % of the annual benthic N remineralisation respectively. Denitrification was dominated by coupled nitrification–denitrification throughout the study. Potential assimilation of nitrogen by benthic microalgae (BMA) accounted for between 1 and 30 % of remineralised N, and was greatest during winter when bottom light was higher. Macrofauna biomass tended to be highest at intermediate benthic respiration rates (?1,000 μmol O₂ m^?2 h^?1), and appeared to become limited as respiration increased above this point. While bioturbation did not significantly affect net fluxes, macrofauna biomass was correlated with increased light rates of NH₄ ⁺ flux which may have masked reductions in NH₄ ⁺ flux associated with BMA assimilation during the light. Peaks in net N₂ fluxes at intermediate respiration rates are suggested to be associated with the stimulation of potential denitrification sites due to bioturbation by burrowing macrofauna. NO₃ ^? fluxes suggest that nitrification was not significantly limited within respiration range measured during this study, however comparisons with other parts of Moreton Bay suggest that limitation of coupled nitrification–denitrification may occur in sub-tropical systems at respiration rates exceeding ?1,500 μmol O₂ m^?2 h^?1.     

Temporal variations in microbenthic metabolism and inorganic nitrogen fluxes in sandy and muddy sediments of a tidally dominated bay in the northern Wadden Sea

Factors controlling seasonal variations in benthic metabolism (O₂ flux) and dissolved inorganic nitrogen (DIN) fluxes were examined during a 12–14 month period at three intertidal Wadden Sea stations. Since the flux measurements were made as small-scale laboratory core incubations, the results are primarily related to the microbenthic community (microalgae, bacteria, micro-, meio- and small macrofauna) and cannot be considered representative of the total benthic community in the Wadden Sea. Furthermore, it has to be emphasized that light intensity during day-time simulations were constant and saturating at all times. Benthic primary production and oxygen uptake appeared to be temperature dependent with a ‘seasonal Q₁₀’ of 1.7–1.8 and 2.7–4.3, respectively. Inundation had no effect on oxygen fluxes as evidenced by similar sediment respiration with and without water cover. A stronger temperature dependence of primary production in muddy than in sandy sediment indicated that the overall control in the latter may be complex due to factors like macrofaunal grazing and nutrient availability. Benthic respiration may not be controlled by temperature alone, as sedimentary organic matter content correlated significantly with both temperature and benthic respiration. Annual gross primary production in high intertidal sandy sediment was 10 and 50% higher than in low intertidal sandy and muddy sediments, respectively. Since annual benthic community respiration was 2 times higher in muddy than sandy sediments, the annual net primary production was about 0 in the former and 17–19 mol C m^?2 yr^?1 in the latter. However, heterotrophic contribution by larger faunal components as well as removal of organic carbon by waves and tidal currents, which are not included here, may balance the budget at the sandy stations. There was no or only weak relationships between (light and dark) DIN exchange and factors like temperature, sedimentary organic content, and oxygen fluxes. Factors related to nutrient fluxes, such as denitrification and nutrient concentration in the overlying water, may have hampered any such relationships. In fact, DIN fluxes at all three stations appeared to be strongly controlled by DIN concentrations in the overlying water. On an annual basis, the sediment appeared to be a net sink for DIN.     

The meagre future of benthic fauna in a coastal sea—Benthic responses to recovery from eutrophication in a changing climate

Nutrient loading and climate change affect coastal ecosystems worldwide. Unravelling the combined effects of these pressures on benthic macrofauna is essential for understanding the future functioning of coastal ecosystems, as it is an important component linking the benthic and pelagic realms. In this study, we extended an existing model of benthic macrofauna coupled with a physical–biogeochemical model of the Baltic Sea to study the combined effects of changing nutrient loads and climate on biomass and metabolism of benthic macrofauna historically and in scenarios for the future. Based on a statistical comparison with a large validation dataset of measured biomasses, the model showed good or reasonable performance across the different basins and depth strata in the model area. In scenarios with decreasing nutrient loads according to the Baltic Sea Action Plan but also with continued recent loads (mean loads 2012–2014), overall macrofaunal biomass and carbon processing were projected to decrease significantly by the end of the century despite improved oxygen conditions at the seafloor. Climate change led to intensified pelagic recycling of primary production and reduced export of particulate organic carbon to the seafloor with negative effects on macrofaunal biomass. In the high nutrient load scenario, representing the highest recorded historical loads, climate change counteracted the effects of increased productivity leading to a hyperbolic response: biomass and carbon processing increased up to mid‐21st century but then decreased, giving almost no net change by the end of the 21st century compared to present. The study shows that benthic responses to environmental change are nonlinear and partly decoupled from pelagic responses and indicates that benthic–pelagic coupling might be weaker in a warmer and less eutrophic sea.     

Spatial and temporal variability in benthic processes along a mangrove-seagrass transect near the Bangrong Mangrove,Thailand

Benthic primary production and nutrient dynamics were examined along a transect in the Bangrong mangrove forest in Thailand. Six stations were established extending from a high-intertidal site within the mangrove forest to low-intertidal flats and seagrass beds in front of the mangrove forest. Benthic processes (O₂ and CO₂ fluxes) and nutrient dynamics (mineralization, sediment-water fluxes, pore water and sediment pools) were measured under light and dark conditions during wet and dry seasons over a 2-yr period. The sediments were mostly autotrophic, only the mangrove forest sites were net heterotrophic during the wet season. Maximum daily net primary production was found at the non-vegetated tidal flats (40–75 mmol O₂ m^-2d^-1), where light and nutrient availability were highest. The variation in benthic mineralization along the transect was minor (1.6–4.3 mmol CO₂ m^-2h^-1) and did not reflect the large changes inorganic matter content (organic carbon: 0.7–4.2% DW) and quality (C:N ratio varied from 25 to 100), suggesting that the mineralizable pool of organic matter was of similar magnitude at all sites. There was only minor seasonal variation in rates of mineralization. The net primary production showed more variation with lower rates in the mangrove forest (reduced with 74%) and higher rates at the tidal flats (increased with 172%) and in the seagrass beds (increased with 228%) during the wet season. The nutrient pools and fluxes across the sediment-water interface were generally low along the transect, and the sediments were efficient in retaining nitrogen in the nutrient limited mangrove/seagrass environment. Pools and fluxes of phosphorus were generally very low suggesting that benthic primary production was phosphorus limited along the transect. This revised version was published online in July 2006 with corrections to the Cover Date.     

Spatial Heterogeneity of Surface CO<Subscript>2</Subscript> Fluxes in a Newly Created Eastmain-1 Reservoir in Northern Quebec,Canada

We assessed the spatial variability in both, surface carbon dioxide (CO₂) fluxes and in pelagic respiration rates in a newly created 600-km² boreal reservoir, located in Northern Quebec. We show that total CO₂ emission to the atmosphere was highest in the first year after flooding, and that surface fluxes were spatially heterogeneous. This spatial heterogeneity was not random, but was linked to the pre-flood landscapes: reservoir areas overlying former peatbogs and mature forests had the highest average emissions, whereas areas overlying former non-forest and burned soils had the lowest emissions. Total reservoir emissions appeared to decline exponentially in the next 2 years, and so did the degree of spatial heterogeneity in surface fluxes, suggesting a progressive weakening of the link to the pre-flood landscapes, and a homogenization of reservoir processes. We show that pelagic respiration rates were also initially high and spatially heterogeneous, the latter linked to the pre-flood landscapes. A simple, first-order mass balance for the first 3 years after flooding was used to derive potential benthic CO₂ production rates, and thus to apportion the total reservoir emissions between its pelagic and benthic components. Extrapolation of the observed declines (normal exponential) in total emission, as well as of the underlying pelagic and benthic fluxes, results in a large underestimation of the fluxes for the fourth year, relative to the measured emissions. We postulate that the initial exponential decline in total emissions is driven primarily by the patterns of decomposition of surface plant biomass, whereas at later stages emission is increasingly dominated by sediment and pelagic respiration, which decline in time at a slower rate.     

Modelling marine ecosystem response to climate change and trawling in the North Sea

The marine ecosystem response to climate change and demersal trawling was investigated using the coupled hydrodynamic-biogeochemical water column model GOTM-ERSEM-BFM for three contrasting sites in the North Sea. Climate change forcing was derived from the HadRM3-PPE-UK regional climate model for the UK for the period 1950–2100 using historical emissions and a medium emissions scenario (SRESA1B). Effects of demersal trawling were implemented as an additional mortality on benthic fauna, and changes in the benthic–pelagic nutrient and carbon fluxes. The main impacts of climate change were (i) a temperature-driven increase in pelagic metabolic rates and nutrient cycling, (ii) an increase in primary production fuelled by recycled nutrients, (iii) a decrease in benthic biomass due to increased benthic metabolic rates and decreased food supply as a result of the increased pelagic cycling, and (iv) a decrease in near-bed oxygen concentrations. The main impacts of trawling were (i) reduced benthic biomass due to the increased mortality, and (ii) the increased benthic–pelagic nutrient fluxes, with these effects counteracting each other, and relatively small changes in other variables. One important consequence was a large decrease in the de-nitrification flux predicted at the two summer-stratified sites because less benthic nitrate was available. The effects of trawling scaled linearly with fishing effort, with greatest sensitivity to fishing in summer compared to fishing in winter. The impacts of climate change and trawling were additive, suggesting little or no non-linear interactions between these disturbances.     

Biogeochemical processes at the sediment–water interface, Bombah Broadwater, Myall Lakes

Myall Lakes has experienced algal blooms in recent years which threaten water quality. Biomarkers, benthic fluxes measured with chambers, and pore water metabolites were used to identify the nature and reactivity of organic matter (OM) in the sediments of Bombah Broadwater (BB), and the processes controlling sediment-nutrient release into the overlying waters. The OM in the sediments was principally from algal sources although terrestrial OM was found near the Myall River. Terrestrial faecal matter was identified in muddy sediments and was probably sourced via runoff from farm lands. The reactive OM which released nutrients into the overlying waters was from diatoms, dinoflagellates and probably cyanobacteria. Microcystis filaments were observed in surface sediments. OM degradation rates varied between 5.3 and 47.1 mmol m^?2 day^?1 (64–565 mg m^?2 day^?1), were highest in the muddy sediments and sulphate reduction rates accounted for 20–40% of the OM degraded. Diatoms, being heavy sink rapidly, and are an important vector to transport catchment N and P to sites of denitrification and P-trapping in the sediments. Denitrification rates (mean ～4 mmol N m^?2 day^?1), up to 7 mmol N m^?2 day^?1 (105 mg N m^?2 day^?1) were measured, and denitrification efficiencies were highest (mean = 86 ± 4%) in the sandy sediments (～20% of the area of BB), but lower in the muddy sediments (mean = 63 ± 15%). These differences probably result from higher OM loads and anaerobic respiration in muddy sediments. Most DIP (>70%) from OM degradation was not released into overlying waters but remained trapped in surface sediments. Biophysical (advective) processes were responsible for the measured metabolite (O₂, CO₂, DSi, DIN and DIP) fluxes across the sediment–water interface.     

Nutrient limitation and algal blooms in urbanizing tidal creeks

Tidal creeks are commonly found in low energy systems on the East and Gulf Coasts of the United States, and are often subject to intense watershed human development. Many of these creeks are receiving urban and suburban runoff containing nutrients, among other pollutants. During the period 1993-2001, we studied three tidal creeks located in southeastern North Carolina, a rapidly urbanizing area. All three creeks received anthropogenic nutrient loading. Oligohaline to mesohaline stations in upper tidal creek regions had much higher nutrient (especially nitrate-N) concentrations than lower creek areas, and hosted spring and summer phytoplankton blooms that at times exceeded 200 μg chlorophyll a l⁻¹. Phytoplankton biomass during winter was low at all stations in all three creeks. Spring and summer nutrient addition bioassay experiments were conducted to characterize the nutrients limiting phytoplankton growth. Water from high salinity stations in all three creeks always showed significant positive responses to nitrate-N inputs, even at concentrations as low as 50 μg N l⁻¹. Low salinity stations in upper creek areas often showed significant responses to nitrate-N inputs, but on occasion showed sensitivity to phosphorus inputs as well, indicating the influence of anthropogenic nitrate loading. During several experiments, one of the upper stations showed no positive response to nutrient inputs, indicating that these stretches were nutrient replete, and further phytoplankton growth appeared to be light-limited either by phytoplankton self-shading or turbidity. Water from upper creek areas yielded much higher chlorophyll a concentrations in bioassay experiments than did lower creek water. In general, these urbanizing tidal creeks were shown to be very sensitive to nitrogen loading, and provide a physical environment conducive to phytoplankton bloom formation in nutrient-enriched areas. Tidal creeks are important ecological resources in that they are considered to be nursery areas for many species of fish and shellfish. To protect the ecological function of these small, but very abundant estuarine systems, management efforts should recognize their susceptibility to algal blooms and focus on control of nonpoint source nutrient inputs, especially nitrogen.