Remote sensing of assimilation number for marine phytoplankton

Estimating primary production at large spatial scales is key to our understanding of the global carbon cycle. Algorithms to estimate primary production are well established and have been used in many studies with success. One of the key parameters in these algorithms is the chlorophyll-normalised production rate under light saturation (referred to as the light saturation parameter or the assimilation number). It is known to depend on temperature, light history and nutrient conditions, but assigning a magnitude to it at particular space-time points is difficult. In this paper, we explore two models to estimate the assimilation number at the global scale from remotely-sensed data that combine methods to estimate the carbon-to-chlorophyll ratio and the maximum growth rate of phytoplankton. The inputs to the algorithms are the surface concentration of chlorophyll, seasurface temperature, photosynthetically-active radiation af the surface of the sea, sea surface nutrient concentration and mixed-layer depth. A large database of in situ estimates of the assimilation number is used to develop the models and provide elements of validation. The comparisons with in situ observations are promising and global maps of assimilation number are produced.

Assimilation of remotely-sensed optical properties to improve marine biogeochemistry modelling

In this paper we evaluate whether the assimilation of remotely-sensed optical data into a marine ecosystem model improves the simulation of biogeochemistry in a shelf sea. A localized Ensemble Kalman filter was used to assimilate weekly diffuse light attenuation coefficient data, Kd(443) from SeaWiFs, into an ecosystem model of the western English Channel. The spatial distributions of (unassimilated) surface chlorophyll from satellite, and a multivariate time series of eighteen biogeochemical and optical variables measured in situ at one long-term monitoring site were used to evaluate the system performance for the year 2006. Assimilation reduced the root mean square error and improved the correlation with the assimilated Kd(443) observations, for both the analysis and, to a lesser extent, the forecast estimates, when compared to the reference model simulation. Improvements in the simulation of (unassimilated) ocean colour chlorophyll were less evident, and in some parts of the Channel the simulation of this data deteriorated. The estimation errors for the (unassimilated) in situ data were reduced for most variables with some exceptions, e.g. dissolved nitrogen. Importantly, the assimilation adjusted the balance of ecosystem processes by shifting the simulated food web towards the microbial loop, thus improving the estimation of some properties, e.g. total particulate carbon. Assimilation of Kd(443) outperformed a comparative chlorophyll assimilation experiment, in both the estimation of ocean colour data and in the simulation of independent in situ data. These results are related to relatively low error in Kd(443) data, and because it is a bulk optical property of marine ecosystems. Assimilation of remotely-sensed optical properties is a promising approach to improve the simulation of biogeochemical and optical variables that are relevant for ecosystem functioning and climate change studies.

Particulate and dissolved aluminum and titanium in the upper water column of the Atlantic Ocean

This study investigates the oceanic behavior of the lithogenic trace elements Al and Ti in the upper 200 m of the Atlantic Ocean. The distribution of both metals in the dissolved and particulate phases was assessed along an E-W transect in the eastern tropical North Atlantic (December 2009) and along a meridional Atlantic transect (April-May 2010). The surface water concentrations of particulate and dissolved Al and Ti reflected the previously observed pattern of atmospheric inputs into the Atlantic Ocean. Subsurface minima at stations with pronounced fluorescence maxima were observed, suggesting a link between biological productivity and the removal of both dissolved and particulate Al and Ti. This may include uptake mechanisms, adsorption and aggregation processes on biogenic particle surfaces and the formation of large, fast sinking biogenic particles, e.g., fecal pellets. Residence times in the upper water column (100 m) of the tropical and subtropical North Atlantic were estimated to range in the order of days to weeks in the particulate phases (Al: 3-22 days, Ti: 4-37 days) and were 0.9-3.8 years for Al and 10-31 years for Ti in the dissolved phases. Longer residence times in both phases in the South Atlantic are consistent with lower biological productivity and decreased removal rates. In the upper water column, Al was predominantly present in the dissolved form, whereas Ti mostly occurred in the particulate form. Largest deviations in the partition coefficients between the particulate and dissolved phases were found in the surface waters, together with excess dissolved Al over Ti compared to the crustal source. This likely reflects elevated dissolution of Al compared to Ti from atmospheric mineral particles.

Combined Effects Of Body Size Food Concentration And Season On Physiology Of Mytilus-Edulis

Multivariate experiments are used to study the effects of body size, food concentration, and season on the oxygen consumption, ammonia excretion, food assimilation efficiency and filtration rate of Mytilus edulis adults. Food concentrations and season affect both the intercept and the slope of the allometric equation describing oxygen uptake as a function of body size. Multiple regression and response surface techniques are used to describe and illustrate the complex relationship between metabolic rate, ration, season and the body size of M. edulis. Filtration rate has a relatively low weight exponent Q> = 038) and the intercept for the allometric equation is not significantly affected by food concentration, season or acclimation temperatures between 5 and 20 °C. Food assimilation efficiency declines exponentially with increasing food concentration and is dependent on body size at high food levels. The rate of ammonia excretion shows a similar seasonal cycle to that of oxygen consumption. They are both minimal in the autumn/winter and reach a maximum in the spring /summer.

Rates Of Nitrogen-Excretion By Species Of Mytilus (Bivalvia - Mollusca)

The results of experiments recorded by Bayne & Scullard (1977) confirmed earlier studies (Bayne, 1973) in describing a decline in the rate of oxygen uptake (Vo2) by Mytilus edulis during starvation, eventually reaching a steady-state value, called the standard rate of oxygen consumption. Earlier experiments had also shown that if such starved mussels were fed, oxygen uptake increased rapidly to a high level called the active rate of oxygen consumption (Thompson & Bayne, 1972; Bayne, Thompson & Widdows, 1973). Some of this increase in metabolic rate is undoubtedly due to an increased filtration rate that is stimulated by the presence of food (the ‘mechanical cost of feeding’ discussed by Bayne et al. 1976), and part is due to the ‘physiological costs of feeding’, which includes energy utilized in digestion and assimilation of the food, and energy that is lost during deamination and other catabolic processes that accompany digestion (Warren & Davis, 1967). Increases in metabolic rate associated with feeding have been called the specific dynamic action (SDA) of the ration (see Harper, 1971, for a discussion) or the apparent SDA (Beamish, 1974)5 and they have been related to aspects of protein metabolism (Krebs, 1964). This paper describes the results of some experiments designed to examine the relationships between SDA and ammonia excretion in Mytilus edulis L.

Apparent Specific Dynamic Action In Mytilus-Edulis L

The results of experiments recorded by Bayne & Scullard (1977) confirmed earlier studies (Bayne, 1973) in describing a decline in the rate of oxygen uptake (Vo2) by Mytilus edulis during starvation, eventually reaching a steady-state value, called the standard rate of oxygen consumption. Earlier experiments had also shown that if such starved mussels were fed, oxygen uptake increased rapidly to a high level called the active rate of oxygen consumption (Thompson & Bayne, 1972; Bayne, Thompson & Widdows, 1973). Some of this increase in metabolic rate is undoubtedly due to an increased filtration rate that is stimulated by the presence of food (the ‘mechanical cost of feeding’ discussed by Bayne et al. 1976), and part is due to the ‘physiological costs of feeding’, which includes energy utilized in digestion and assimilation of the food, and energy that is lost during deamination and other catabolic processes that accompany digestion (Warren & Davis, 1967). Increases in metabolic rate associated with feeding have been called the specific dynamic action (SDA) of the ration (see Harper, 1971, for a discussion) or the apparent SDA (Beamish, 1974)5 and they have been related to aspects of protein metabolism (Krebs, 1964). This paper describes the results of some experiments designed to examine the relationships between SDA and ammonia excretion in Mytilus edulis L.

Fine scale variability in methanol uptake and oxidation in the micro-layer and near-surface waters of the Atlantic

The aim of this research was to make the first depth profiles of the microbial assimilation of methanol carbon and its oxidation to carbon dioxide and use as an energy source from the microlayer to 1000 m. Some of the highest reported methanol oxidation rate constants of 0.5–0.6 d−1 were occasionally found in the microlayer and immediately underlying waters (10 cm depth), albeit these samples also showed the greatest heterogeneity compared to other depths down to 1000 m. Methanol uptake into the particulate phase was exceptionally low in microlayer samples, suggesting that any methanol utilised by microbes in this environment is for energy generation. The sea surface microlayer and 10 cm depth also showed a higher proportion of bacteria with a low DNA content, and bacterial leucine uptake rates in surface microlayer samples were either less than or the same as those in the underlying 10 cm layer. The average methanol oxidation and particulate rates were however statistically the same throughout the depths sampled, although the latter were highly variable in the near-surface 0.25–2 m compared to deeper depths. The statistically significant relationship demonstrated between uptake of methanol into particles and bacterial leucine incorporation suggests that many heterotrophic bacteria could be using methanol carbon for cellular growth. On average, methanol bacterial growth efficiency (BGEm) in the top 25 m of the water column is 6% and decreases with depth. Although, for microlayer and 10 cm-depth samples, BGEm is less than the near-surface 25–217 cm, possibly reflecting increased environmental UV stress resulting in increased maintenance costs, i.e. energy required for survival. We conclude that microbial methanol uptake rates, i.e. loss from seawater, are highly variable, particularly close to the seawater surface, which could significantly impact upon seawater concentrations and hence the air–sea flux.

Fine-scale variability in methanol uptake and oxidation: from the microlayer to 1000 m.

The aim of this research was to make the first depth profiles of the microbial assimilation of methanol carbon and its oxidation to carbon dioxide and use as an energy source from the microlayer to 1000 m. Some of the highest reported methanol oxidation rate constants of 0.5–0.6 d−1 were occasionally found in the microlayer and immediately underlying waters (10 cm depth), albeit these samples also showed the greatest heterogeneity compared to other depths down to 1000 m. Methanol uptake into the particulate phase was exceptionally low in microlayer samples, suggesting that any methanol utilised by microbes in this environment is for energy generation. The sea surface microlayer and 10 cm depth also showed a higher proportion of bacteria with a low DNA content, and bacterial leucine uptake rates in surface microlayer samples were either less than or the same as those in the underlying 10 cm layer. The average methanol oxidation and particulate rates were however statistically the same throughout the depths sampled, although the latter were highly variable in the near-surface 0.25–2 m compared to deeper depths. The statistically significant relationship demonstrated between uptake of methanol into particles and bacterial leucine incorporation suggests that many heterotrophic bacteria could be using methanol carbon for cellular growth. On average, methanol bacterial growth efficiency (BGEm) in the top 25 m of the water column is 6% and decreases with depth. Although, for microlayer and 10 cm-depth samples, BGEm is less than the near-surface 25–217 cm, possibly reflecting increased environmental UV stress resulting in increased maintenance costs, i.e. energy required for survival. We conclude that microbial methanol uptake rates, i.e. loss from seawater, are highly variable, particularly close to the seawater surface, which could significantly impact upon seawater concentrations and hence the air–sea flux.

Low colloidal associations of aluminium and titanium in surface waters of the tropical Atlantic

The distribution of dissolved, soluble and colloidal fractions of Al and Ti was assessed by ultrafiltration studies in the upper water column of the eastern tropical North Atlantic. The dissolved fractions of both metals were found to be dominated by the soluble phase smaller than 10 kDa. The colloidal associations were very low (0.2–3.4%) for Al and not detectable for Ti. These findings are in some contrast to previous estimations for Ti and to the predominant occurrence of both metals as hydrolyzed species in seawater. However, low tendencies to form inorganic colloids can be expected, as in seawater dissolved Al and dissolved Ti are present within their inorganic solubility levels. In addition, association with functional organic groups in the colloidal phase is unlikely for both metals. Vertical distributions of the dissolved fractions showed surface maxima with up to 43 nM of Al and 157 pM of Ti, reflecting their predominant supply from atmospheric sources to the open ocean. In the surface waters, excess dissolved Al over dissolved Ti was present compared to the crustal source, indicating higher solubility and thus elevated inputs of dissolved Al from atmospheric mineral particles. At most stations, subsurface minima of Al and Ti were observed and can be ascribed to scavenging processes and/or biological uptake. The dissolved Al concentrations decreased by 80–90% from the surface maximum to the subsurface minimum. Estimated residence times in the upper 100 m of the water column ranged between 1.6 and 4 years for dissolved Al and between 14 and 17 years for dissolved Ti. The short residence times are in some contrast to the low colloidal associations of Al and Ti and the assumed role of colloids as intermediates in scavenging processes. This suggests that either the removal of both metals occurs predominantly via direct transfer of the hydrolyzed species into the particulate fraction or that the colloidal phase is rapidly turned over in the upper water column.

Parameterisation of bivalve functional traits for mechanistic eco-physiological dynamic energy budget (DEB) models

Mechanistic models such as those based on dynamic energy budget (DEB) theory are emergent ecomechanics tools to investigate the extent of fitness in organisms through changes in life history traits as explained by bioenergetic principles. The rapid growth in interest around this approach originates from the mechanistic characteristics of DEB, which are based on a number of rules dictating the use of mass and energy flow through organisms. One apparent bottleneck in DEB applications comes from the estimations of DEB parameters which are based on mathematical and statistical methods (covariation method). The parameterisation process begins with the knowledge of some functional traits of a target organism (e. g. embryo, sexual maturity and ultimate body size, feeding and assimilation rates, maintenance costs), identified from the literature or laboratory experiments. However, considering the prominent role of the mechanistic approach in ecology, the reduction of possible uncertainties is an important objective. We propose a revaluation of the laboratory procedures commonly used in ecological studies to estimate DEB parameters in marine bivalves. Our experimental organism was Brachidontes pharaonis. We supported our proposal with a validation exercise which compared life history traits as obtained by DEBs (implemented with parameters obtained using classical laboratory methods) with the actual set of species traits obtained in the field. Correspondence between the 2 approaches was very high (>95%) with respect to estimating both size and fitness. Our results demonstrate a good agreement between field data and model output for the effect of temperature and food density on age-size curve, maximum body size and total gamete production per life span. The mechanistic approach is a promising method of providing accurate predictions in a world that is under in creasing anthropogenic pressure.

The influence of ocean acidification on nitrogen regeneration and nitrous oxide production in the North-West European shelf sea

The assimilation and regeneration of dissolved inorganic nitrogen, and the concentration of N2O, was investigated at stations located in the NW European shelf sea during June/July 2011. These observational measurements within the photic zone demonstrated the simultaneous regeneration and assimilation of NH4+, NO2− and NO3−. NH4+ was assimilated at 1.82–49.12 nmol N L−1 h−1 and regenerated at 3.46–14.60 nmol N L−1 h−1; NO2− was assimilated at 0–2.08 nmol N L−1 h−1 and regenerated at 0.01–1.85 nmol N L−1 h−1; NO3− was assimilated at 0.67–18.75 nmol N L−1 h−1 and regenerated at 0.05–28.97 nmol N L−1 h−1. Observations implied that these processes were closely coupled at the regional scale and nitrogen recycling played an important role in sustaining phytoplankton growth during the summer. The [N2O], measured in water column profiles, was 10.13 ± 1.11 nmol L−1 and did not strongly diverge from atmospheric equilibrium indicating that sampled marine regions where neither a strong source nor sink of N2O to the atmosphere. Multivariate analysis of data describing water column biogeochemistry and its links to N-cycling activity failed to explain the observed variance in rates of N-regeneration and N-assimilation, possibly due to the limited number of process rate observations. In the surface waters of 5 further stations, Ocean Acidification (OA) bioassay experiments were conducted to investigate the response of NH4+ oxidising and regenerating organisms to simulated OA conditions, including the implications for [N2O]. Multivariate analysis was undertaken which considered the complete bioassay dataset of measured variables describing changes in N-regeneration rate, [N2O] and the biogeochemical composition of seawater. While anticipating biogeochemical differences between locations, we aimed to test the hypothesis that the underlying mechanism through which pelagic N-regeneration responded to simulated OA conditions was independent of location and that a mechanistic understanding of how NH4+ oxidation, NH4+ regeneration and N2O production responded to OA could be developed. Results indicated that N-regeneration process responses to OA treatments were location specific; no mechanistic understanding of how N-regeneration processes respond to OA in the surface ocean of the NW European shelf sea could be developed.

Investigating the inter-relationships between water attenuated irradiance, primary production and DMS(P)

Both solar irradiance and primary production have been proposed as independent controls on seawater dimethyl sulphide (DMS) and dimethylsulphoniopropionate (DMSP) concentrations. However, irradiance also drives photosynthesis, and thus influences a complex set of inter-related processes that modulate marine DMS. We investigate the potential inter-relationships between the rate of primary production (carbon assimilation), water-attenuated irradiance and DMS/DMSP dynamics by applying correlation analysis to a high resolution, concurrently sampled in situ data set from a range of latitudes covering multiple biogeochemical provinces from 3 of the 4 Longhurst biogeochemical domains. The combination of primary production (PP) and underwater irradiance (Iz) within a multivariate regression model is able to explain 55% of the variance in DMS concentrations from all depths within the euphotic zone and 66% of the variance in surface DMS concentrations. Contrary to some previous studies we find a variable representing biological processes is necessary to better account for the variance in DMS. We find that the inclusion of Iz accounts for variance in DMS that is independent from the variance explained by PP. This suggests an important role for solar irradiance (beyond the influence of irradiance upon primary production) in mediating the relationship between the productivity of the ecosystem, DMS/DMSP production and ambient seawater DMS concentrations.

Bridging the gap between marine biogeochemical and fisheries sciences; configuring the zooplankton link

Exploring climate and anthropogenic impacts on marine ecosystems requires an understanding of how trophic components interact. However, integrative end-to-end ecosystem studies (experimental and/or modelling) are rare. Experimental investigations often concentrate on a particular group or individual species within a trophic level, while tropho-dynamic field studies typically employ either a bottom-up approach concentrating on the phytoplankton community or a top-down approach concentrating on the fish community. Likewise the emphasis within modelling studies is usually placed upon phytoplankton-dominated biogeochemistry or on aspects of fisheries regulation. In consequence the roles of zooplankton communities (protists and metazoans) linking phytoplankton and fish communities are typically under-represented if not (especially in fisheries models) ignored. Where represented in ecosystem models, zooplankton are usually incorporated in an extremely simplistic fashion, using empirical descriptions merging various interacting physiological functions governing zooplankton growth and development, and thence ignoring physiological feedback mechanisms. Here we demonstrate, within a modelled plankton food-web system, how trophic dynamics are sensitive to small changes in parameter values describing zooplankton vital rates and thus the importance of using appropriate zooplankton descriptors. Through a comprehensive review, we reveal the mismatch between empirical understanding and modelling activities identifying important issues that warrant further experimental and modelling investigation. These include: food selectivity, kinetics of prey consumption and interactions with assimilation and growth, form of voided material, mortality rates at different age-stages relative to prior nutrient history. In particular there is a need for dynamic data series in which predator and prey of known nutrient history are studied interacting under varied pH and temperature regimes.

The influence of ocean acidification on nitrogen regeneration and nitrous oxide production in the North-West European shelf sea

The assimilation and regeneration of dissolved inorganic nitrogen, and the concentration of N2O, was investigated at stations located in the NW European shelf sea during June/July 2011. These observational measurements within the photic zone demonstrated the simultaneous regeneration and assimilation of NH4+, NO2− and NO3−. NH4+ was assimilated at 1.82–49.12 nmol N L−1 h−1 and regenerated at 3.46–14.60 nmol N L−1 h−1; NO2− was assimilated at 0–2.08 nmol N L−1 h−1 and regenerated at 0.01–1.85 nmol N L−1 h−1; NO3− was assimilated at 0.67–18.75 nmol N L−1 h−1 and regenerated at 0.05–28.97 nmol N L−1 h−1. Observations implied that these processes were closely coupled at the regional scale and nitrogen recycling played an important role in sustaining phytoplankton growth during the summer. The [N2O], measured in water column profiles, was 10.13 ± 1.11 nmol L−1 and did not strongly diverge from atmospheric equilibrium indicating that sampled marine regions where neither a strong source nor sink of N2O to the atmosphere. Multivariate analysis of data describing water column biogeochemistry and its links to N-cycling activity failed to explain the observed variance in rates of N-regeneration and N-assimilation, possibly due to the limited number of process rate observations. In the surface waters of 5 further stations, Ocean Acidification (OA) bioassay experiments were conducted to investigate the response of NH4+ oxidising and regenerating organisms to simulated OA conditions, including the implications for [N2O]. Multivariate analysis was undertaken which considered the complete bioassay dataset of measured variables describing changes in N-regeneration rate, [N2O] and the biogeochemical composition of seawater. While anticipating biogeochemical differences between locations, we aimed to test the hypothesis that the underlying mechanism through which pelagic N-regeneration responded to simulated OA conditions was independent of location and that a mechanistic understanding of how NH4+ oxidation, NH4+ regeneration and N2O production responded to OA could be developed. Results indicated that N-regeneration process responses to OA treatments were location specific; no mechanistic understanding of how N-regeneration processes respond to OA in the surface ocean of the NW European shelf sea could be developed.

The influence of ocean acidification on nitrogen regeneration and nitrous oxide production in the northwest European shelf sea

The assimilation and regeneration of dissolved inorganic nitrogen, and the concentration of N₂O, was investigated at stations located in the NW European shelf sea during June/July 2011. These observational measurements within the photic zone demonstrated the simultaneous regeneration and assimilation of NH₄⁺, NO₂⁻ and NO₃⁻. NH₄⁺ was assimilated at 1.82–49.12 nmol N L⁻¹ h⁻¹ and regenerated at 3.46–14.60 nmol N L⁻¹ h⁻¹; NO₂^- was assimilated at 0–2.08 nmol N L⁻¹ h⁻¹ and regenerated at 0.01–1.85 nmol N L⁻¹ h⁻¹; NO₃⁻ was assimilated at 0.67–18.75 nmol N L⁻¹ h⁻¹ and regenerated at 0.05–28.97 nmol N L⁻¹ h⁻¹. Observations implied that these processes were closely coupled at the regional scale and that nitrogen recycling played an important role in sustaining phytoplankton growth during the summer. The [N₂O], measured in water column profiles, was 10.13 ± 1.11 nmol L⁻¹ and did not strongly diverge from atmospheric equilibrium indicating that sampled marine regions were neither a strong source nor sink of N₂O to the atmosphere. Multivariate analysis of data describing water column biogeochemistry and its links to N-cycling activity failed to explain the observed variance in rates of N-regeneration and N-assimilation, possibly due to the limited number of process rate observations. In the surface waters of five further stations, ocean acidification (OA) bioassay experiments were conducted to investigate the response of NH₄⁺ oxidising and regenerating organisms to simulated OA conditions, including the implications for [N₂O]. Multivariate analysis was undertaken which considered the complete bioassay data set of measured variables describing changes in N-regeneration rate, [N₂O] and the biogeochemical composition of seawater. While anticipating biogeochemical differences between locations, we aimed to test the hypothesis that the underlying mechanism through which pelagic N-regeneration responded to simulated OA conditions was independent of location. Our objective was to develop a mechanistic understanding of how NH₄⁺ regeneration, NH₄⁺ oxidation and N₂O production responded to OA. Results indicated that N-regeneration process responses to OA treatments were location specific; no mechanistic understanding of how N-regeneration processes respond to OA in the surface ocean of the NW European shelf sea could be developed.