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

Technical note: Estimating light-use efficiency of benthic habitats using underwater O2 eddy covariance

Light-use efficiency defines the ability of primary producers to convert sunlight energy to primary production and is computed as the ratio between the gross primary production and the intercepted photosynthetic active radiation. While this measure has been applied broadly within terrestrial ecology to investigate habitat resource-use efficiency, it remains underused within the aquatic realm. This report provides a conceptual framework to compute hourly and daily light-use efficiency using underwater O-2 eddy covariance, a recent technological development that produces habitat-scale rates of primary production under unaltered in situ conditions. The analysis, tested on two benthic flux datasets, documents that hourly light-use efficiency may approach the theoretical limit of 0.125 O-2 per photon under low-light conditions, but it decreases rapidly towards the middle of the day and is typically 10-fold lower on a 24 h basis. Overall, light- use efficiency provides a useful measure of habitat functioning and facilitates site comparison in time and space.Peer reviewe

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

Seasonal rates of benthic primary production in a Greenland fjord measured by aquatic eddy correlation

We present the first year-round estimates of benthic primary production at four contrasting shallow (3–22 m depth) benthic habitats in a southwest Greenland fjord. In situ measurements were performed using the noninvasive aquatic eddy-correlation (EC) oxygen (O2) flux method. A series of high-quality multiple-day EC data sets document the presence of a year-round productive benthic phototrophic community. The shallow-water sites were on average autotrophic during the spring and summer months, up to 43.6 mmol O2 m22 d21, and heterotrophic or close to metabolic balance during the autumn and winter. Substantial benthic gross primary production (GPP) was measured year-round. The highest GPP rates were measured during the spring, up to 5.7 mmol O2 m22 h21 (136.8 mmol O2 m22 d21), and even at low light levels (, 80 mmol quanta m22 s21) during late autumn and winter we measured rates of up to 1.8 mmol O2 m22 h21 (43.2 mmol O2 m22 d21) during peak irradiance. The benthic phototrophic communities responded seasonally to ambient light levels and exhibited year-round high photosynthetic efficiency. In situ downwelling irradiances as low as , 2 mmol quanta m22 s21 induced an autotrophic response and light saturation indices (Ik) were as low as 11 mmol quanta m22 s21 in the winter. On an annual timescale, the average areal rate of benthic GPP was 11.5 mol O2 m22 yr21, which is , 1.4 times higher than the integrated gross pelagic primary production of the , 30–50 m deep photic zone of the fjord. These results document the importance of benthic photosynthesis on an ecosystem level and indicate that the benthic phototrophic compartment should be accounted for when assessing carbon and nutrient budgets as well as responses of coastal Arctic ecosystems to climate change

Benthic microalgal production in the Arctic: Applied methods and status of the current database

The current database on benthic microalgal production in Arctic waters comprises 10 peer-reviewed and three unpublished studies. Here, we compile and discuss these datasets, along with the applied measurement approaches used. The latter is essential for robust comparative analysis and to clarify the often very confusing terminology in the existing literature. Our compilation demonstrates that i) benthic microalgae contribute significantly to coastal ecosystem production in the Arctic, and ii) benthic microalgal production on average exceeds pelagic productivity by a factor of 1.5 for water depths down to 30 m. We have established relationships between irradiance, water depth and benthic microalgal productivity that can be used to extrapolate results from quantitative experimental studies to the entire Arctic region. Two different approaches estimated that current benthic microalgal production in the Arctic is between 1.1 and 1.6×107 tons C year-1. Climate change is expected to increase the overall primary production and affect the balance between pelagic and benthic productivity in the Arctic. It is therefore imperative to get better quantitative understanding of the relationship between increased freshwater run-off, shrinking sea-ice cover, light availability and benthic primary production to assess future impact on the Arctic food web and trophic coupling. © 2009 by Walter de Gruyter

Microbial degradation dynamics of farmed kelp deposits from Saccharina latissima and Alaria esculenta

Seaweed farming is a growing industry worldwide, and its sustainable management requires detailed knowledge about the environmental implications of detrital release. This study investigates benthic degradation of kelp detritus in defaunated mesocosms. The degradation dynamics were investigated over several weeks by resolving O2 and dissolved inorganic carbon (DIC) fluxes as a function of detritus amendments (0.15 g wet weight [WW] m-2 to 1 kg WW m-2), temperature (8 and 15°C), and presence of O2 for 2 commercially important kelp species: Saccharina latissima and Alaria esculenta. Kelp fragments were deposited in 2 different ways to simulate oxic and anoxic degradation: on the sediment surface (surface amendments) and just below the oxic surface sediment layer (subsurface amendments). All amendments resulted in high initial O2 consumption followed by an exponential decrease in O2 uptake over time. The degradation rates increased linearly with the amount of kelp added for both species and for both types of amendments. S. latissima expressed higher decay constants across all experiments and had a higher percentage turnover of carbon. In some instances, microbial priming apparently enabled enhanced degradation of pre-existing resilient sedimentary carbon. The absolute degradation rates of kelp were reduced in the absence of O2, and sulfate reduction resulted in gradual accumulation of iron sulfide. Lower ambient temperature reduced the benthic mineralization rate of both kelp species, particularly during the initial incubation stages. The current study demonstrates the importance of key variables for microbial kelp degradation in marine sediments and their dynamics—variables that should be carefully considered when assessing environmental implications of seaweed farming.publishedVersio

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

Copepod carcasses as microbial hot spots for pelagic denitrification

Copepods are exposed to a high non-predatory mortality and their decomposing carcasses act as microniches with intensified microbial activity. Sinking carcasses could thereby represent anoxic microenvironment sustaining anaerobic microbial pathways in otherwise oxic water columns. Using non-invasive O-2 imaging, we document that carcasses of Calanus finmarchicus had an anoxic interior even at fully air-saturated ambient O-2 level. The extent of anoxia gradually expanded with decreasing ambient O-2 levels. Concurrent microbial sampling showed the expression of nitrite reductase genes (nirS) in all investigated carcass samples and thereby documented the potential for microbial denitrification in carcasses. The nirS gene was occasionally expressed in live copepods, but not as consistently as in carcasses. Incubations of sinking carcasses in (15)NO3-amended seawater demonstrated denitrification, of which on average 34%+/- 17% (n=28) was sustained by nitrification. However, the activity was highly variable and was strongly dependent on the ambient O-2 levels. While denitrification was present even at air-saturation (302 mol L-1), the average carcass specific activity increased several orders of magnitude to approximate to 1 nmol d(-1) at 20% air-saturation (55 mol O-2 L-1) at an ambient temperature of 7 degrees C. Sinking carcasses of C. finmarchicus therefore represent hotspots of pelagic denitrification, but the quantitative importance as a sink for bioavailable nitrogen is strongly dependent on the ambient O-2 level. The importance of carcass associated denitrification could be highly significant in O-2 depleted environments such as Oxygen Minimum Zones (OMZ)