Manganese and Iron Oxidation During Benthic Oxygenic Photosynthesis

The effect of benthic oxygenic photosynthesis on sediment-water fluxes of manganese and iron was studied for an intertidal sediment. Undisturbed sediments were incubated at an incident surface irradiance of 250 μE m⁻² s⁻¹at 26 °C. Oxygenic photosynthesis was selectively inhibited by adding [3-(3,4-dichloro)-1,1-dimethyl-urea] (DCMU). Benthic fluxes were determined experimentally from the change in manganese and iron concentrations in the overlying water, and were predicted from the pore water concentration gradients at the sediment-water interface assuming molecular diffusion as the transport mechanism. The experimental fluxes of manganese and iron in DCMU-treated cores amounted to −0·84 and −0·59 mmol m⁻²day⁻¹, respectively, and were directed from the sediment towards the overlying water. In the control cores, showing high rates of benthic oxygenic photosynthesis, the fluxes of manganese and iron were directed towards the sediment, 0·06 and 0·01 mmol m⁻²day⁻¹, respectively. Mass balances for the 0·1–0·14 cm thick oxic zone, calculated from the experimental fluxes and the predicted fluxes, suggest a minimum areal reoxidation of 0·6 mmol m⁻²day⁻¹for manganese and of 0·48 mmol m⁻²day⁻¹for iron in cores showing benthic photosynthesis. The estimated turnover times for dissolved Mn²⁺and dissolved Fe²⁺in the oxic surface layer during benthic photosynthesis were 0·8 and 0·25 h, respectively. Sediment oxygen microprofiles and the sediment pH profiles suggest that chemical precipitation and reoxidation dominates the retention of manganese and iron during benthic oxygenic photosynthesis in shallow intertidal sediments.     

Methane sources,sinks and fluxes in a temperate tidal Lagoon: The Arcachon lagoon (SW France)

The Arcachon lagoon is a 156 km² temperate mesotidal lagoon dominated by tidal flats (66% of the surface area). The methane (CH₄) sources, sinks and fluxes were estimated from water and pore water concentrations, from chamber flux measurements at the sediment–air (low tide), sediment–water and water–air (high tide) interfaces, and from potential oxidation and production rate measurements in sediments. CH₄ concentrations in waters were maximal (500–1000 nmol l⁻¹) in river waters and in tidal creeks at low tide, and minimal in the lagoon at high tide (<50 nmol l⁻¹). The major CH₄ sources are continental waters and the tidal pumping of sediment pore waters at low tide. Methanogenesis occurred in the tidal flat sediments, in which pore water concentrations were relatively high (2.5–8.0 μmol l⁻¹). Nevertheless, the sediment was a minor CH₄ source for the water column and the atmosphere because of a high degree of anaerobic and aerobic CH₄ oxidation in sediments. Atmospheric CH₄ fluxes at high and low tide were low compared to freshwater wetlands. Temperate tidal lagoons appear to be very minor contributor of CH₄ to global atmosphere and to open ocean.     

Trophic importance of diatoms in an intertidal Zostera noltii seagrass bed: Evidence from stable isotope and fatty acid analyses

A current predominant paradigm emphasizes the role of epiphytic algae for invertebrates in most seagrass food webs. However, in some intertidal Zostera noltii beds, epiphyte biomass is very low compared to microphytobenthos and seagrass biomasses. We assessed the role of microphytobenthos in a temperate intertidal Z. noltii bed by combining stable isotope and fatty acid (FA) analyses on primary producers, composite sources — suspended particulate organic matter (SPOM) and sediment surface organic matter (SSOM) — and the main macrofaunal consumers. Z. noltii showed high δ¹³C (−9.9‰) and high 18:2(n-6) and 18:3(n-3) contents. Microphytobenthos was slightly more ¹³C-depleted (−15.4‰) and had high levels of diatom markers: 14:0, 16:1(n-7)c, 20:5(n-3). Low mean δ¹³C (−22.0‰) and large amounts of diatom and bacteria (18:1(n-7)c) markers indicated that SPOM was mainly composed of a mixture of fresh and decayed pelagic diatoms. Higher mean δ¹³C (−17.9‰) and high amounts of diatom FAs were found in SSOM, showing that microphytobenthic diatoms dominate. Very low percentages of 18:2(n-6) and 18:3(n-3) in consumers indicated a low contribution of Z. noltii material to their diets. Grazers, deposit and suspension-deposit feeders had δ¹³C close to microphytobenthos and high levels of diatom FAs, confirming that microphytobenthos represented the main part of their diet. Lower δ¹³C and higher amounts of flagellate FAs – 22:6(n-3) and 16:4(n-3) – in suspension feeders indicated that their diet resulted from a mixture of SPOM and microphytobenthos. These results demonstrate that invertebrates do not consume high amounts of seagrass and highlight the main role of benthic diatoms in this intertidal seagrass bed.     

Nutrient exchange across the sediment-water interface in the Potomac River estuary

The flux of ammonia, phosphate, silica and radon-222 from Potomac tidal river and estuary sediments is controlled by processes occurring at the sediment-water interface and within surficial sediment. Calculated diffusive fluxes range between 0·6 and 6·5 mmol m^?2 day^?1 for ammonia, 0·020 and 0·30 mmol m^?2 day^?1 for phosphate, and 1·3 and 3·8 mmol m^?2 day^?1 for silica. Measured in situ fluxes range between 1 and 21 mmol m^?2 day^?1 for ammonia, 0·1 and 2·0 mmol m^?2 day^?1 for phosphate, and 2 and 19 mmol m^?2 day^?1 for silica. The ratio of in situ fluxes to diffusive fluxes (flux enhancement) varied between 1·6 and 5·2 in the tidal river, between 2·0 and 20 in the transition zone, and from 1·3 to 5·1 in the lower estuary. The large flux enhancements from transition zone sediments are attributed to macrofaunal irrigation. Nutrient flux enhancements are correlated with radon flux enhancements, suggesting that fluxes may originate from a common region and that nutrients are regenerated within the upper 10–20 cm of the sediment column.The low fluxes of phosphate from tidal viver sediments reflect the control benthic sediment exerts on phosphorus through sorption by sedimentary iron oxyhydroxides. In the tidal river, benthic fluxes of ammonia and phosphate equal one-half and one-third of the nutrient input of the Blue Plains sewage treatment plant. In the tidal Potomac River, benthic sediment regeneration supplies a significant fraction of the nutrients utilized by primary producers in the water column during the summer months.     

The effect of tidal range on the flushing of ammonium from intertidal sediments of the Tagus estuary,Portugal

The time-course evolution of ammonium concentration has been examined in the flood water during the first 25 min of tidal inundation. The way this transport fluctuates with the tidal ranges and wind conditions was investigated. Flood water was collected at three sites, located along a transect from the lower to the upper intertidal area of the Tagus estuary. At spring and intermediate tides, the periods of air exposure vary slightly along the transect due to the high tidal amplitude and the flatness of the area, but the upper site remains uncovered at neap tide over the entire tidal cycle. At each site, sampling was performed at different tidal ranges covering the neap-spring tidal cycle and wind conditions. Ammonium was determined in the flood water at short time intervals: 1, 2, 3, 4, 5, 10, 15, 20 and 25 min. A clear pattern was observed along the transect: considerable quantities of ammonium were exported from the sediment to the water column at the beginning of the inundation, ranging from 0.2 to 4.8 mmol m⁻² d⁻¹. The highest transport was recorded at the lower intertidal site under spring tide conditions, which corresponds to the higher energetic situation and shorter emersion period. The lowest transport was observed at the upper intertidal site during the first inundation that followed three days of neap tide and continuous exposure of the sediment to the air. The value rates (0.2–4.8 mmol m⁻² d⁻¹) were one order of magnitude higher than those calculated from molecular diffusion (0.07 – 0.16 mmol m⁻² d⁻¹). This study points to the importance of the tidal flushing of ammonium from the intertidal sediments, and its spatial and tidal fluctuation.     

Benthic nutrient fluxes along the Laurentian Channel: Impacts on the N budget of the St. Lawrence marine system

Water column concentrations and benthic fluxes of dissolved inorganic nitrogen (DIN) and oxygen (DO) were measured in the Gulf of St. Lawrence and the Upper and Lower St. Lawrence Estuary (USLE and LSLE, respectively) to assess the nitrogen (N) budget in the St. Lawrence (SL) system, as well as to elucidate the impact of bottom water hypoxia on fixed-N removal in the LSLE. A severe nitrate deficit, with respect to ambient phosphate concentrations (N*∼−10 μmol L⁻¹), was observed within and in the vicinity of the hypoxic bottom water of the LSLE. Given that DO concentrations in the water column have remained above 50 μmol L⁻¹, nitrate reduction in suboxic sediments, rather than in the water column, is most likely responsible for the removal of fixed N from the SL system. Net nitrate fluxes into the sediments, derived from pore water nitrate concentration gradients, ranged from 190 μmol m⁻² d⁻¹ in the hypoxic western LSLE to 100 μmol m⁻² d⁻¹ in the Gulf. The average total benthic nitrate reduction rate for the Laurentian Channel (LC) is on the order of 690 μmol m⁻² d⁻¹, with coupled nitrification-nitrate reduction accounting for more than 70%. Using average nitrate reduction rates derived from the observed water column nitrate deficit, the annual fixed-N elimination within the three main channels of the Gulf of St. Lawrence and LSLE was estimated at 411 × 10⁶ t N, yielding an almost balanced N budget for the SL marine system.     

The Role of Microphytobenthic Primary Production in a Mediterranean Mussel Culture Area

The production and biomass of microphytobenthos in a Mediterranean mussel farm was studied during 1991–92. Gross and net microphytobenthic production and respiration were calculated from oxygen fluxes in transparent and black bell jars at two stations; sediments under a mussel table and reference sediments, both located at 5 m depth. Net oxygen fluxes were mainly negative under the mussel tables (average −19·5 mg O₂ m⁻² h⁻¹, CV=132%), and microphytobenthos production could not meet the sediment oxygen demand; in the reference sediments, microphytobenthos production was responsible for net oxygen production (average +13·0 mg O₂ m⁻² h⁻¹, CV=118%). Benthic respiration rates were, on average, 47·3 mg O₂ m⁻² h⁻¹(CV=82%) under the tables and 27·7 mg O₂ m⁻² h⁻¹(CV=45%) in reference sediments. Aerobic respiration could remineralize less than 2% of the biodeposited carbon under the tables, implying that a large amount of organic material is accumulating under the tables, and that most of the degradation will be anaerobic. Gross microbenthic production showed sharp changes between 1991 and 1992 under the mussel tables and for reference sediments (averages 20·98 mg O₂ m⁻² h⁻¹, CV=135% and 33 mg O₂ m⁻² h⁻¹, CV=48%, respectively). Despite the negative oxygen balance in the sediments under the tables, microphytobenthos was more productive than phytoplankton in bottom waters. Per unit area, phytoplankton was more productive than microphytobenthos at both stations, especially in the area of the mussel tables, where phytoplanktonic production was enhanced by the excretion products of mussels. Microphytobenthos was composed mainly of diatoms in the sediments under the tables, while in reference sediments, the population was more diverse, with algae containing chlorophyllbalso present. Chlorophyllaconcentration in sediments under the tables was 207 mg m⁻²(CV=73%) and 95 mg m⁻²(CV=28%) in reference sediments; the stock of plant pigments was increased under the tables by biodeposition. Microphytobenthos constitutes a compartment with an important contribution in biomass, but also in oxygen production.     

Diurnal heterogeneity in silicic acid fluxes in shallow coastal sites: Causes and implications

In shallow coastal areas the amplitude and range of benthic silicic acid fluxes can have a significant influence on benthic–pelagic coupling and the functioning of the pelagic system. To explore the oscillation in fluxes over the diurnal cycle and in particular the influence of microphytobenthos (MPB), an experiment was carried out in a shallow subtidal site in the Bay of Brest (France). Benthic chambers were employed over a 48 h period to measure the variability in silicic acid and oxygen fluxes; MPB migration was investigated using a diving Pulse Amplitude Modulated (PAM) fluorometer and uptake rhythms of silicic acid by natural MPB populations were measured using the ³²Si isotope. It was discovered that silicic acid fluxes fluctuated greatly throughout the diurnal period resulting in an oscillation in the availability of this nutrient for phytoplankton communities. The uptake of silicic acid by the MPB was quantified for the first time and highlighted a 2-fold increase in the demand from night to afternoon periods. The combined silicic acid uptake and the concentration of cells at the sediment–water interface, forming a dense biofilm of MPB, were postulated to be the main processes reducing effluxes at midday. Our work highlighted the many processes which influence silicic acid effluxes in shallow coastal areas and the possible interaction between uptake and dissolution processes. The variations in benthic fluxes over the diurnal period were comparable to observations reported at the seasonal scale. Therefore, up-scaling hourly flux observations to daily and annual estimates should be undertaken with caution. Further we suggest that the main processes influencing flux oscillations over the diurnal period should be considered when planning sampling strategies and extrapolating to larger time scales.     

Observations of resuspended diatoms in the turbid tidal edge

Observations of resuspended diatoms in the shallow waters (<60 cm) of the turbid tidal edge are described for single sites on two tidal flats–the Molenplaat in the Westerschelde estuary, and the Hond in the Ems-Dollard estuary, The Netherlands. High concentrations of chlorophyll-a (chl-a) were observed in the trailing edge of the ebbing tide in water depths of <20 cm, after which concentrations decreased markedly. Peak mean values were 19 μg chl-a l⁻¹ in 10 cm of water at the Molenplaat, and 45 μg chl-a l⁻¹ in 5 cm of water at the Hond. Similar trends were observed on the flooding tide, although peak values were far less pronounced (6 and 30 μg chl-a l⁻¹ respectively). Microscopic examination of the diatom community within the turbid tidal edge at the Molenplaat revealed that peaks in biomass were caused by suspended benthic diatoms, as well as the large centric diatom Coscinodiscus sp., particularly on the ebb tide. Planktonic diatoms other than Coscinodiscus sp. were more randomly distributed and did not appear to follow any particular trend. It would seem that as the tide recedes, resuspended benthic diatoms and large Coscinodiscus sp. cells become concentrated in the shallow water. However, the virtual absence of Coscinodiscus sp. from the leading edge of the flooding tide suggests that most of the resuspended cells do not settle to the seabed, but are washed away into the channels. The small peak of benthic diatoms at the leading edge of the flood tide is most likely resuspended locally from the sediment, along with large numbers of diatom frustules.     

Annual Variation of Benthic Nutrient Fluxes in Shallow Coastal Waters (Gulf of Trieste, Northern Adriatic Sea)

Abstract. Benthic fluxes of dissolved N. Si and P nutrients, alkalinity, dissolved inorganic C (DIC), and O₂ from sediments in the Gulf of Trieste (northern Adriatic, Italy) were measured monthly for 16 months, using laboratory incubated flux chambers at in siru temperatures in the dark. The annual average fluxes were: 02 = -19.3 ± 8.2, DIC = 13.7 ± 9.6, NO3 = -0.04 ± 0.16, NH4 = 0.3 ± 0.4. PO₄= 4.001 ± 0.01, Si = 0.9 ± 0.1 mmol m-² d^-1, with strong temporal fluctuations. The highest effluxes of all nutrients and DIC were observed in the summer. Small effluxes of DIC and NH4 and influxes of Si and PO4 were observed in late winter. Only NH₄ (ca. 50%) and Si (ca. 70%) fluxes were significantly correlated with temperature. This correlation suggests that the rate of downward input and the quality of sedimented organic matter (autochthonous and allochthonous) were superimposed on the temperature fluctuations. High DIC, NH₄ and Si effluxes observed in May 1993 during low temperature were due to the degradation of sedimentary organic matter produced by an early spring bloom of benthic microalgae which occurred about 6 weeks earlies while the autumn phytoplankton bloom was simultaneously reflected in enhanced benthic fluxes due to higher temperature. The role of benthic biological advection in this transport across the sediment-water interface, evaluated by comparison between measured benthic and calculated diffusive fluxes from nutrient pore water concentrations, was of minor importance. This is probably due to low infaunal activity throughout the year it was localized mostly in the narrow surficial layer. The annual average diffusive fluxes of NH4 and PO4 were higher than those measured, probably due to the presence of nitrificationdenitrifi-cation processes and redox-dependent chemical reactions at the oxic sediment-water interface, respectively. Only during bottom-water hypoxia in September 1993 did strong PO₄ effluxes prevail. Calculations based on the Redfield stoichiometry of oxic decomposition of organic N to NH4 and NO3, and differences between diffusive and measured NH₄ fluxes showed that denitrifkation averaged 0.8 mmol m^-2 d^-1. Significant correlations between NH₄ and PO₄ DIC and Si, and NH4 and Si fluxes suggested their parallel regeneration and utilization at the sediment-water interface. The nutrient fluxes observed were not significantly linked to O₂ consumption, suggesting also that anaerobic oxidation processes were important at the sediment-water interface in the gulf. The N, P and Si nutriqnts released from sediment pore waters are probably utilized in benthic microalgal and bottorn-hater primary production. This indicates that pelagic and benthic communities in the central part of the Gulf of Trieste function relatively independently of each other.     

Potentially mobile pools of phosphorus and silicon in sediment from the Bay of Brest: Interactions and implications for phosphorus dynamics

Competitive interactions between silicate and phosphate at ligand exchange sites in the sediment surface layer may increase the release of phosphorus (P) from the sediment into the water column. In this study, the role of silicon (Si) in the release of P from the sediment surface layer was studied in a marine estuarine environment, the Bay of Brest, with the aid of a sequential sediment fractionation procedure developed for P, and the addition of inorganic or diatom-bound Si to surface sediment samples in vitro. The potentially mobile pools of P in the surface sediment (loosely bound P + Fe/Al-bound-P) amounted to 5.0 μmol g⁻¹ dry sed., 42% of the total extractable and 33% of the total amount of P in the sediment, while the similarly extracted pools of Si were bigger (ca. 20 μmol g⁻¹ dry sed., 50% of the total extractable Si). Additions of inorganic Si increased the concentration of dissolved P in the sediment interstitial water in a bottle experiment, and the addition of both inorganic Si and cultivated diatoms to intact sediment cores increased the outward flux of dissolved P. Model calculations based on the regression equation from the bottle experiment and Si and P water column data showed that the sedimentation of spring diatoms could cause Si pulses to the sediment which would produce a P flux to the water column of ca. 44 μmol m⁻² d⁻¹. Field data from the bay show that in spring, decreases in P and Si and an increase in chl a due to diatom production are often followed by a small separate P peak which may be caused by Si-induced P fluxes from the sediment surface.     

Submarine groundwater discharge (SGD) as a main nutrient source for benthic and water-column primary production in a large intertidal environment of the Yellow Sea

The biogeochemistry and magnitude of submarine groundwater discharge (SGD) was investigated in one of the largest tidal flat ecosystems worldwide, along the Yellow Sea coast. A representative semi-enclosed embayment located in the south eastern Yellow Sea, Hampyeong Bay, was chosen for this purpose. Groundwater and seawater samples were collected in three seasons (May, July, and September) and analyzed for Ra isotopes, nutrients, and photosynthetic pigments. The biogeochemistry of SGD was strongly influenced by tidal oscillations and seasonal precipitation changes and switched from a brackish, nutrient-enriched regime in May and July to an exclusively saline regime, with lower nutrient concentrations, in September. SGD magnitudes, calculated by using a ²²⁶Ra mass balance model, were 0.14 m³ m^? 2 d^? 1 in May and 0.35 m³ m^? 2 d^? 1 in September. A nutrient mass balance was established for the two campaigns, which suggests that SGD causes the flushing of substantial amounts of pore water nutrients into this embayment; because of SGD, the embayment acts as a source of dissolved inorganic silicates (DSi) that are transported to the open ocean. Potential C fixation rates derived from this nutrient mass balance were compared with two different models for water-column phytoplankton productivity based on water-column Chl a and local irradiation levels. The Chl a-based models generally showed lower C fixation rates than the nutrient-based mass balance, indicating removal of up to 70% of the nutrients by other primary producers, such as benthic algae. During monsoon season, when benthic algal biomass is high and nutrient fluxes are substantial due to a terrestrial component, SGD — driven benthic primary production could play a significant role in this large tidal flat ecosystem.     

Microphytobenthic biomass and species composition in intertidal flats of the Nakdong River estuary, Korea

Intertidal microphytobenthos (MPB) were investigated monthly from August 2006 to March 2008 at four different sites in the sand flats of Nakdong River estuary, Korea. Samples of surface sediment (ca. 1 cm) were collected, and chlorophyll a was extracted as biomass estimation. Species identification and enumeration were carried out by light microscopy, assisted where necessary by scanning electronic microscopy. Biomass varied between 0.47 and 16.58 μg cm⁻³, abundance changed from 5.25 to 414.75 × 10³ cells cm⁻³, while the Shannon diversity indexes ranged between 0.69 and 2.35 H′. Thirty-nine MPB taxa were identified, primarily composed of epipelic diatoms, among which Amphora and Navicula were the most abundant genera. Based on the biomass, abundance, species composition and their dynamics, MPB assemblages of sampling sites were grouped into three distinct communities corresponding to their sediment composition characteristics. Multivariate correlation analysis revealed that biomass was positively related to mud and very fine sand, negatively related to fine and medium sand, but not significantly related to environmental factors such as pore water nutrients, light intensity and salinity, which fluctuated rapidly during emersion period. Cluster analysis corroborated the division of MPB communities according to site types on seasonal scales, and also showed seasonality between sites by cluster of all summer groups. Principal component analysis identified that variability in species composition was significantly affected by mud, very fine sand, fine sand, light intensity, and sediment temperature. This study suggests that sediment composition plays an important role in the functioning of intertidal MPB communities in estuarine ecosystems.     

Benthic fluxes in a tropical Estuary and their role in the ecosystem

In-situ measurements of benthic fluxes of oxygen and nutrients were made in the subtidal region of the Mandovi estuary during premonsoon and monsoon seasons to understand the role of sediment–water exchange processes in the estuarine ecosystem. The Mandovi estuary is a shallow, highly dynamic, macrotidal estuary which experiences marine condition in the premonsoon season and nearly fresh water condition in the monsoon season. The benthic flux of nutrients exhibited strong seasonality, being higher in the premonsoon compared to the monsoon season which explains the higher ecosystem productivity in the dry season in spite of negligible riverine nutrient input. NH₄⁺ was the major form of released N comprising 70–100% of DIN flux. The benthic respiration rate varied from −98.91 to −35.13 mmol m⁻² d⁻¹, NH₄⁺ flux from 5.15 to 0.836 mmol m⁻² d⁻¹, NO₃⁻ + NO₂⁻ from 0.06 to −1.06 mmol m⁻² d⁻¹, DIP from 0.12 to 0.23 mmol m⁻² d⁻¹ and SiO₄⁴⁻ from 5.78 to 0.41 mmol m⁻² d⁻¹ between premonsoon to monsoon period. The estuarine sediment acted as a net source of DIN in the premonsoon season, but changed to a net sink in the monsoon season. Variation in salinity seemed to control NH₄⁺ flux considerably. Macrofaunal activities, especially bioturbation, enhanced the fluxes 2–25 times. The estuarine sediment was observed to be a huge reservoir of NH₄⁺, PO₄³⁻ and SiO₄⁴⁻ and acted as a net sink of combined N because of the high rate of benthic denitrification as it could remove 22% of riverine DIN influx thereby protecting the eco system from eutrophication and consequent degradation. The estuarine sediment was responsible for ∼30–50% of the total community respiration in the estuary. The benthic supply of DIN, PO₄³⁻ and SiO₄⁴⁻ can potentially meet 49%, 25% and 55% of algal N, P and Si demand, respectively, in the estuary. Based on these observations we hypothesize that it is mainly benthic NH₄⁺ efflux that sustains high estuarine productivity in the NO₃⁻ depleted dry season.     

The cycling and oxidation pathways of organic carbon in a shallow estuary along the Texas Gulf Coast

The cycling and oxidation pathways of organic carbon were investigated at a single shallow water estuarine site in Trinity Bay, Texas, the uppermost lobe of Galveston Bay, during November 2000. Radio-isotopes were used to estimate sediment mixing and accumulation rates, and benthic chamber and pore water measurements were used to determine sediment-water exchange fluxes of oxygen, nutrients and metals, and infer carbon oxidation rates. Using ⁷Be and ²³⁴Th_XS, the sediment-mixing coefficient (D_b) was 4.3 ± 1.8 cm² y⁻¹, a value that lies at the lower limit for marine environments, indicating that mixing was not important in these sediments at this time. Sediment accumulation rates (S_a), estimated using ¹³⁷Cs and ²¹⁰Pb_XS, were 0.16 ± 0.02 g cm⁻² y⁻¹. The supply rate of organic carbon to the sediment-water interface was 30 ± 3.9 mmol C m⁻² d⁻¹, of which ∼10% or 2.9 ± 0.44 mmol C m⁻² d⁻¹was lost from the system through burial below the 1-cm thick surface mixed layer. Measured fluxes of O₂ were 26 ± 3.8 mmol m⁻² d⁻¹ and equated to a carbon oxidation rate of 20 ± 3.3 mmol C m⁻² d⁻¹, which is an upper limit due to the potential for oxidation of additional reduced species. Using organic carbon gradients in the surface mixed layer, carbon oxidation was estimated at 2.6 ± 1.1 mmol C m⁻² d⁻¹. Independent estimates made using pore water concentration gradients of ammonium and C:N stoichiometry, equaled 2.8 ± 0.46 mmol C m⁻² d⁻¹. The flux of DOC out of the sediments (DOC_efflux) was 5.6 ± 1.3 mmol C m⁻² d⁻¹. In general, while mass balance was achieved indicating the sediments were at steady state during this time, changes in environmental conditions within the bay and the surrounding area, mean this conclusion might not always hold. These results show that the majority of carbon oxidation occurred at the sediment-water interface, via O₂ reduction. This likely results from the high frequency of sediment resuspension events combined with the shallow sediment mixing zone, leaving anaerobic oxidants responsible for only ∼10–15% of the carbon oxidized in these sediments.     

Benthic photosynthesis in submerged Wadden Sea intertidal flats

In this study we compare benthic photosynthesis during inundation in coarse sand, fine sand, and mixed sediment (sand/mud) intertidal flats in the German Wadden Sea. In situ determinations of oxygen-, DIC- and nutrient fluxes in stirred benthic chamber incubations were combined with measurements of sedimentary chlorophyll, incident light intensity at the sediment surface and scalar irradiance within the sediment. During submergence, microphytobenthos was light limited at all study sites as indicated by rapid response of gross photosynthesis to increasing incident light at the sea floor. However, depth integrated scalar irradiance was 2 to 3 times higher in the sands than in the mud. Consequently, gross photosynthesis in the net autotrophic fine sand and coarse sand flats during inundation was on average 4 and 11 times higher than in the net heterotrophic mud flat, despite higher total chlorophyll concentration in mud. Benthic photosynthesis may be enhanced in intertidal sands during inundation due to: (1) higher light availability to the microphytobenthos in the sands compared to muds, (2) more efficient transport of photosynthesis-limiting solutes to the microalgae with pore water flows in the permeable sands, and (3) more active metabolic state and different life strategies of microphytobenthos inhabiting sands.     

Distribution and contribution of major phytoplankton groups to carbon cycling across contrasting conditions of the subtropical northeast Atlantic Ocean

The relation between trophic regime and phytoplankton composition and function in oceanic systems is well accepted in oceanography. However, the relative dynamics and carbon cycling contributions of different phytoplankton groups across gradients of ocean richness are not fully understood. In this work we investigated phytoplankton dynamics along two transects from the NW African coastal upwelling to open-ocean waters of the north Atlantic subtropical gyre. We adopted a pigment-based approach to characterize community structure and to quantify group-specific growth and grazing rates and associated carbon fluxes. Changes in pigment cell concentration during the incubation experiments due to photoadaptation were corrected to obtain reliable rates. The oceanic region was dominated by Prochlorococcus (PRO) (45±7% of total chlorophyll a) while diatoms dominated in upwelling waters (40±37%). Phytoplankton grew faster (μ=0.78±0.26 d⁻¹) and free of nutrient limitation (μ/μ_n=0.98±0.42) in the coastal upwelling region, with all groups growing at similar rates. In oceanic waters, the growth rate of bulk phytoplankton was lower (μ=0.52±0.16 d⁻¹) and nutrient limited (μ/μ_n=0.68±0.19 d⁻¹). Diatoms (0.80±0.39 d⁻¹) and Synechococcus (SYN) (0.72±0.25 d⁻¹) grew faster than Prymnesiophyceae (PRYMN) (0.62±0.26 d⁻¹) and PRO (0.46±0.18 d⁻¹). The growth rates of PRO and SYN were moderately nutrient limited (μ/μ_n=0.81 and 0.91, respectively), while the limitation for diatoms (μ/μ_n=0.71) and PRYMN (μ/μ_n=0.37) was more severe. Microzooplankton grazing rate was higher in upwelling (0.68±0.32 d⁻¹) than in oceanic waters (0.37±0.19 d⁻¹), but represented the main loss pathway for phytoplankton in both systems (m/μ=0.90±0.32 and 0.69±0.24, respectively). Carbon flux through phytoplankton, produced and grazed, increased from offshore to coastal (∼2 to ∼200 μg C L⁻¹ d⁻¹), with diatoms dominating the flux in the upwelling region (52%) while PRYMN (40%) and PRO (30%) dominated in the open ocean.     

Swash zone sediment fluxes: Field observations

This paper describes newly obtained, high-frequency observations of beach face morphological change over numerous tidal cycles on a macrotidal sandy beach made using a large array of ultrasonic altimeters. These measurements enable the net cross-shore sediment fluxes associated with many thousands of individual swash events to be quantified. It is revealed that regardless of the direction of net morphological change on a tidal time scale, measured net fluxes per event are essentially normally distributed, with nearly equal numbers of onshore and offshore-directed events. The majority of swash events cause net cross-shore sediment fluxes smaller than ± 50 kg m^− 1 and the mean sediment flux per swash event is only O(± 1 kg m^− 1) leading to limited overall morphological change. However, much larger events which deposit or remove hundreds of kilograms of sand per meter width of beach occur at irregular intervals throughout the course of a tide. It was found that swash–swash interactions tend to increase the transport potential of a swash event and the majority of the swash events that cause these larger values of sediment flux include one or more interactions. The majority of the larger sediment fluxes were therefore measured in the lower swash zone, close to the surf/swash boundary where swash–swash interactions are most common. Despite the existence of individual swash events that can cause fluxes of sediment that are comparable to those observed on a tidal time scale, frequent reversals in transport direction act to limit net transport such that the beach face volume remains in a state of dynamic equilibrium and does not rapidly erode or accrete.     

Temporal variation in the vertical distribution of microphytobenthos in intertidal flats of the Nakdong River estuary,Korea

The vertical distribution of microphytobenthos (MPB) was investigated in intertidal flats of the Nakdong River estuary over different temporal periods. The MPB biomass were measured as chlorophyll a concentration in the sediment, and monitored at two different depth scales. From August 2006 to August 2007, monthly sampling was carried out at four sites to a depth of 10 cm. The vertical distribution exhibited an exponential decline with depth and the slope was closely related to sediment composition and MPB biomass in the surface sediment. The two slightly muddy sand (5–25% mud, <63 μm) sites presented similar trends of seasonal variation to a sediment depth of 10 cm, and no statistical difference was observed in their initial values and the slope of the exponential curves applied to each data set. The site with higher mud content also had higher surface biomass, and the fitted exponential curve had a larger slope than sites with more sand content. The seasonal variations of phaeo-pigments compared to chlorophyll a revealed higher grazing stress on the MPB in summer at the three slightly muddy sand sites, and low grazing stress all year around at the sand (<5% mud) site. In January, April and October 2007, and January 2008, diurnal sampling was conducted at hourly intervals at one slightly muddy sand site. The investigation in January 2007 which was conducted at 1 cm intervals to a depth of 10 cm sediment showed the migration mainly existed in the upper 1 cm. The other diurnal sampling was undertaken to a 1 cm depth at 0.5 mm intervals. An evident migratory rhythm was present, with migration up to the surface during daytime emersion and down into deeper sediment during night and high tide submersion. Migrating cells could reach sediments as deep as 6 or even 10 mm. When the visible thick biofilms formed, the migratory rhythm changed by keeping higher biomass at the surface even at night and during high tide submersion. Generally, the MPB surface biomass was an important factor in structuring the MPB vertical distribution. The vertical distribution to depths of 10 cm was closely related to the sediment composition over the long term, while the MPB migration influenced the vertical distribution over the short term in the top 1 cm of the sediment.     

Erodibility of a mixed mudflat dominated by microphytobenthos and Cerastoderma edule, East Frisian Wadden Sea,Germany

Sediment erodibility and a range of physical and biological parameters were measured at an intertidal site in the German Wadden Sea area in June, September and November 2002 and February and April 2003 in order to examine the influence of macrozoobenthos and microphytobenthos on sediment erodibility and the temporal variation. The study site was a mixed mudflat situated in the mesotidal Baltrum–Langeoog tidal basin at the East Frisian barrier coast. The mud content at the site was about 35% and the filter-feeding cockle Cerastoderma edule was the dominating macrozoobenthic species (by biomass). The erodibility of the sediment showed strong temporal variation with high erosion thresholds in spring and late summer and significantly lower thresholds during the rest of the study period. The erosion thresholds were strongly dependent on the contents of chlorophyll a (chl a) and colloidal carbohydrates, both indicators of the content of microphytobenthos, in this environment primarily benthic diatoms. The content of microphytobenthos was high in September 2002 and April 2003, and regression analysis indicated that this was the only likely reason for the low erodibility found at these times. A biostabilisation index of about 4.5 was found for a situation with both abundant biofilms and cockles.