An in situ instrument for planar O2 optode measurements at benthic interfaces

A new in situ instrument for two‐dimensional mapping of oxygen in coastal sediments is presented. The measuring principle is described, and potential mechanical disturbances, solute and particle smearing associated with the measurements, and calibration routines are evaluated. The first in situ measurements obtained in two different benthic communities are presented. In a shallow photosynthetic sediment (1 m of water depth), an extensive horizontal and temporal variation in the O2 distribution caused by benthic photosynthesis and irrigating fauna was resolved. Repetitive planar optode measurements performed along a transect in central Øresund, Denmark (17 m of water depth) revealed a positive correlation between the apparent O2 penetration depths (OP) measured with a lateral distance <5.0 mm, whereas OP measured with a larger horizontal distance (up to 50 m) were not correlated. Consequently, the OP varied in patches with a characteristic size of 5.0 mm. The instrument described is a powerful new tool for in situ characterization of spatiotemporal variations in O2 distributions within benthic communities. The instrument can be adapted for use at full ocean depths, e.g., on deep‐sea landers or remote operating vehicles

Benthic oxygen exchange in a live coralline algal bed and an adjacent sandy habitat: an eddy covariance study

Coralline algal (maerl) beds are widespread, slow-growing, structurally complex perennial habitats that support high biodiversity, yet are significantly understudied compared to seagrass beds or kelp forests. We present the first eddy covariance (EC) study on a live maerl bed, assessing the community benthic gross primary productivity (GPP), respiration (R), and net ecosystem metabolism (NEM) derived from diel EC time series collected during 5 seasonal measurement campaigns in temperate Loch Sween, Scotland. Measurements were also carried out at an adjacent (~20 m distant) permeable sandy habitat. The O2 exchange rate was highly dynamic, driven by light availability and the ambient tidally-driven flow velocity. Linear relationships between the EC O2 fluxes and available light indicate that the benthic phototrophic communities were lightlimited. Compensation irradiance (Ec) varied seasonally and was typically ~1.8-fold lower at the maerl bed compared to the sand. Substantial GPP was evident at both sites; however, the maerl bed and the sand habitat were net heterotrophic during each sampling campaign. Additional inputs of ~4 and ~7 mol m-2 yr-1 of carbon at the maerl bed and sand site, respectively, were required to sustain the benthic O2 demand. Thus, the 2 benthic habitats efficiently entrap organic carbon and are sinks of organic material in the coastal zone. Parallel deployment of 0.1 m2 benthic chambers during nighttime revealed O2 uptake rates that varied by up to ~8-fold between replicate chambers (from -0.4 to -3.0 mmol O2 m-2 h-1; n = 4). However, despite extensive O2 flux variability on meter horizontal scales, mean rates of O2 uptake as resolved in parallel by chambers and EC were typically within 20% of one another

Estimating respiration rates and secondary production of macrobenthic communities across coastal habitats with contrasting structural biodiversity

A central goal of benthic ecology is to describe the pathways and quantities of energy and material flow in seafloor communities over different spatial and temporal scales. We examined the relative macrobenthic contribution to the seafloor metabolism by estimating respiration and secondary production based on seasonal measurements of macrofauna biomass across key coastal habitats of the Baltic Sea archipelago. Then, we compared the macrofauna estimates with estimates of overall seafloor gross primary production and respiration obtained from the same habitats using the aquatic eddy covariance technique. Estimates of macrobenthic respiration rates suggest habitat-specific macrofauna contribution (%) to the overall seafloor respiration ranked as follows: blue mussel reef (44.5) > seagrass meadow (25.6) > mixed meadow (24.1) > bare sand (17.8) > Fucus-bed (11.1). In terms of secondary production (g C m−2 y−1), our estimates suggest ranking of habitat value as follows: blue mussel reef (493.4) > seagrass meadow (278.5) > Fucus-bed (102.2) > mixed meadow (94.2) > bare sand (52.1). Our results suggest that approximately 12 and 10% of the overall soft-sediment metabolism translated into macrofauna respiration and secondary production, respectively. The hard-bottoms exemplified two end-points of the coastal metabolism, with the Fucus-bed as a high producer and active exporter of organic C (that is, net autotrophy), and the mussel reef as a high consumer and active recycler of organic C (that is, net heterotrophy). Using a combination of metrics of ecosystem functioning, such as respiration rates and secondary production, in combination with direct habitat-scale measurements of O2 fluxes, our study provides a quantitative assessment of the role of macrofauna for ecosystem functioning across heterogeneous coastal seascapes.Peer reviewe

Intracellular nitrate of marine diatoms as a driver of anaerobic nitrogen cycling in sinking aggregates

Diatom-bacteria aggregates are key for the vertical transport of organic carbon in the ocean. Sinking aggregates also represent pelagic microniches with intensified microbial activity, oxygen depletion in the center, and anaerobic nitrogen cycling. Since some of the aggregate-forming diatom species store nitrate intracellularly, we explored the fate of intracellular nitrate and its availability for microbial metabolism within anoxic diatom-bacteria aggregates. The ubiquitous nitrate-storing diatom Skeletonema marinoi was studied as both axenic cultures and laboratory-produced diatom-bacteria aggregates. Stable 15N isotope incubations under dark and anoxic conditions revealed that axenic S. marinoi is able to reduce intracellular nitrate to ammonium that is immediately excreted by the cells. When exposed to a light:dark cycle and oxic conditions, S. marinoi stored nitrate intracellularly in concentrations > 60 mmol L-1 both as free-living cells and associated to aggregates. Intracellular nitrate concentrations exceeded extracellular concentrations by three orders of magnitude. Intracellular nitrate was used up within 2-3 days after shifting diatom-bacteria aggregates to dark and anoxic conditions. Thirty-one percent of the diatom-derived nitrate was converted to nitrogen gas, indicating that a substantial fraction of the intracellular nitrate pool of S. marinoi becomes available to the aggregate-associated bacterial community. Only 5% of the intracellular nitrate was reduced to ammonium, while 59% was recovered as nitrite. Hence, aggregate-associated diatoms accumulate nitrate from the surrounding water and sustain complex nitrogen transformations, including loss of fixed nitrogen, in anoxic, pelagic microniches. Additionally, it may be expected that intracellular nitrate not converted before the aggregates have settled onto the seafloor could fuel benthic nitrogen transformations

Aquatic Eddy Correlation: Quantifying the Artificial Flux Caused by Stirring-Sensitive O2 Sensors

In the last decade, the aquatic eddy correlation (EC) technique has proven to be a powerful approach for non-invasive measurements of oxygen fluxes across the sediment water interface. Fundamental to the EC approach is the correlation of turbulent velocity and oxygen concentration fluctuations measured with high frequencies in the same sampling volume. Oxygen concentrations are commonly measured with fast responding electrochemical microsensors. However, due to their own oxygen consumption, electrochemical microsensors are sensitive to changes of the diffusive boundary layer surrounding the probe and thus to changes in the ambient flow velocity. The so-called stirring sensitivity of microsensors constitutes an inherent correlation of flow velocity and oxygen sensing and thus an artificial flux which can confound the benthic flux determination. To assess the artificial flux we measured the correlation between the turbulent flow velocity and the signal of oxygen microsensors in a sealed annular flume without any oxygen sinks and sources. Experiments revealed significant correlations, even for sensors designed to have low stirring sensitivities of ~0.7%. The artificial fluxes depended on ambient flow conditions and, counter intuitively, increased at higher velocities because of the nonlinear contribution of turbulent velocity fluctuations. The measured artificial fluxes ranged from 2-70 mmol m(-2) d(-1) for weak and very strong turbulent flow, respectively. Further, the stirring sensitivity depended on the sensor orientation towards the flow. For a sensor orientation typically used in field studies, the artificial flux could be predicted using a simplified mathematical model. Optical microsensors (optodes) that should not exhibit a stirring sensitivity were tested in parallel and did not show any significant correlation between O2 signals and turbulent flow. In conclusion, EC data obtained with electrochemical sensors can be affected by artificial flux and we recommend using optical microsensors in future EC-studies

Sediment accumulation and carbon burial in four hadal trench systems

Hadal trenches are considered to act as depocenters for organic material, although pathways for the material transport and deposition rates are poorly constrained. Here we assess focusing, deposition and accumulation of material and organic carbon in four hadal trench systems underlying different surface ocean productivities; the eutrophic Atacama and Kuril-Kamchatka trenches, the mesotrophic Kermadec trench and the oligotrophic Mariana Trench. The study is based on the distributions of naturally occurring 210Pbex, 137Cs and total organic carbon from recovered sediment cores and by applying previously quantified benthic mineralization rates. Periods of steady deposition and discreet mass-wasting deposits were identified from the profiles and the latter were associated with historic recorded seismic events in the respective regions. During periods without mass wasting, the estimated focusing factors along trench axes were elevated, suggesting more or less continuous downslope focusing of material toward the interior of the trenches. The estimated organic carbon deposition rates during these periods exhibited extensive site-specific variability, but were generally similar to values encountered at much shallower settings such as continental slopes and margins. Organic carbon deposition rates during periods of steady deposition were not mirrored by surface ocean productivity, but appeared confounded by local bathymetry. The inclusion of deposition mediated by mass-wasting events enhanced the sediment and organic carbon accumulations for the past ∼ 150 years by up to a factor of ∼ 4. Thus, due to intensified downslope material focusing and infrequent mass-wasting events, hadal trenches are important sites for deposition and sequestration of organic carbon in the deep sea