Increased carbon dioxide availability alters phytoplankton stoichiometry and affects carbon cycling and growth of a marine planktonic herbivore

Rising levels of CO2 in the atmosphere have led to increased CO2 concentrations in the oceans. This enhanced carbon availability to the marine primary producers has the potential to change their nutrient stoichiometry, and higher carbon to nutrient ratios are expected. As a result, the quality of the primary producers as food for herbivores may change. Here, we present experimental work showing the effect of feeding Rhodomonas salina grown under different pCO2 (200, 400 and 800 µatm) on the copepod Acartia tonsa. The rate of development of copepodites decreased with increasing CO2 availability to the algae. The surplus carbon in the algae was excreted by the copepods, with younger stages (copepodites) excreting most of their surplus carbon through respiration, and adult copepods excreting surplus carbon mostly as DOC. We consider the possible consequences of different excretory pathways for the ecosystem. A continued increase in the CO2 availability for primary production, together with changes in the nutrient loading of coastal ecosystems, may cause changes in the trophic links between primary producers and herbivores.

Model results of sensitivity experiments for marine nitrogen cycle in four NetCDF files

The marine nitrogen (N) inventory is thought to be stabilized by negative feedback mechanisms that reduce N inventory excursions relative to the more slowly overturning phosphorus inventory. Using a global biogeochemical ocean circulation model we show that negative feedbacks stabilizing the N inventory cannot persist if a close spatial association of N2 fixation and denitrification occurs. In our idealized model experiments, nitrogen deficient waters, generated by denitrification, stimulate local N2 fixation activity. But, because of stoichiometric constraints, the denitrification of newly fixed nitrogen leads to a net loss of N. This can enhance the N deficit, thereby triggering additional fixation in a vicious cycle, ultimately leading to a runaway N loss. To break this vicious cycle, and allow for stabilizing negative feedbacks to occur, inputs of new N need to be spatially decoupled from denitrification. Our idealized model experiments suggest that factors such as iron limitation or dissolved organic matter cycling can promote such decoupling and allow for negative feedbacks that stabilize the N inventory. Conversely, close spatial co-location of N2 fixation and denitrification could lead to net N loss.

Mineral nitrogen from KCl extractions of soil from the Jena Experiment (Main Experiment up to 15cm depth, year 2008)

As an estimate of plant-available N, this data set contains measurements of inorganic nitrogen (NO3-N and NH4-N, the sum of which is termed mineral N or Nmin) determined by extraction with 1 M KCl solution of soil samples from the main experiment plots of a large grassland biodiversity experiment (the Jena Experiment; see further details below). In the main experiment, 82 grassland plots of 20 x 20 m were established from a pool of 60 species belonging to four functional groups (grasses, legumes, tall and small herbs). In May 2002, varying numbers of plant species from this species pool were sown into the plots to create a gradient of plant species richness (1, 2, 4, 8, 16 and 60 species) and functional richness (1, 2, 3, 4 functional groups). Plots were maintained by bi-annual weeding and mowing. Soil sampling and analysis: Five soil cores (diameter 0.01 m) were taken at a depth of 0 to 0.15 m of the mineral soil from each of the experimental plots in March and October 2008. In October 2008, also the plots of the management experiment, that altered mowing frequency and fertilized subplots (see further details below) were sampled. Samples of the soil cores per plot (subplots in case of the management experiment) were pooled during each sampling campaign. NO3-N and NH4-N concentrations were determined by extraction of soil samples with 1 M KCl solution and were measured in the soil extract with a Continuous Flow Analyzer (CFA, AutoAnalyzer, Seal, Burgess Hill, United Kingdom).

Mineral nitrogen from KCl extractions of soil from the Jena Experiment (Main Experiment up to 15cm depth, year 2006)

As an estimate of plant-available N, this data set contains measurements of inorganic nitrogen (NO3-N and NH4-N, the sum of which is termed mineral N or Nmin) determined by extraction with 1 M KCl solution of soil samples from the main experiment plots of a large grassland biodiversity experiment (the Jena Experiment; see further details below). In the main experiment, 82 grassland plots of 20 x 20 m were established from a pool of 60 species belonging to four functional groups (grasses, legumes, tall and small herbs). In May 2002, varying numbers of plant species from this species pool were sown into the plots to create a gradient of plant species richness (1, 2, 4, 8, 16 and 60 species) and functional richness (1, 2, 3, 4 functional groups). Plots were maintained by bi-annual weeding and mowing. Soil sampling and analysis: Five soil cores (diameter 0.01 m) were taken at a depth of 0 to 0.15 m of the mineral soil from each of the experimental plots in March 2006. In October 2006 also the plots of the management experiment, that altered mowing frequency and fertilized subplots (see further details below) were sampled. Measurements from the management experiment are separated into 0 to 0.08 m and 0.08 to 0.15 m. Samples of the soil cores per plot (subplots in case of the management experiment) were pooled during each sampling campaign. NO3-N and NH4-N concentrations were determined by extraction of soil samples with 1 M KCl solution and were measured in the soil extract with a Continuous Flow Analyzer (CFA, AutoAnalyzer, Seal, Burgess Hill, United Kingdom).

Mineral nitrogen from KCl extractions of soil from the Jena Experiment (Main Experiment up to 15cm depth, year 2007)

As an estimate of plant-available N, this data set contains measurements of inorganic nitrogen (NO3-N and NH4-N, the sum of which is termed mineral N or Nmin) determined by extraction with 1 M KCl solution of soil samples from the main experiment plots of a large grassland biodiversity experiment (the Jena Experiment; see further details below). In the main experiment, 82 grassland plots of 20 x 20 m were established from a pool of 60 species belonging to four functional groups (grasses, legumes, tall and small herbs). In May 2002, varying numbers of plant species from this species pool were sown into the plots to create a gradient of plant species richness (1, 2, 4, 8, 16 and 60 species) and functional richness (1, 2, 3, 4 functional groups). Plots were maintained by bi-annual weeding and mowing. Soil sampling and analysis: Five soil cores (diameter 0.01 m) were taken at a depth of 0 to 0.15 m of the mineral soil from each of the experimental plots in March and October 2007. In March and in October 2007 also the plots of the management experiment, that altered mowing frequency and fertilized subplots (see further details below) were sampled. Samples of the soil cores per plot (subplots in case of the management experiment) were pooled during each sampling campaign. NO3-N and NH4-N concentrations were determined by extraction of soil samples with 1 M KCl solution and were measured in the soil extract with a Continuous Flow Analyzer (CFA, AutoAnalyzer, Seal, Burgess Hill, United Kingdom).

Nitrogen isotopes of bulk sediments for OAE2 samples

Sediment records of the stable isotopic composition of N (d15N) show light d15N values at several sites in the proto-North Atlantic during Oceanic Anoxic Event 2 (OAE 2) at the Cenomanian-Turonian transition (~94 Ma). The low d15N during the event is generally attributed to an increase in N2-fixation and incomplete uptake of ammonium for phytoplankton growth. A compilation of all reliable data for the proto North-Atlantic during OAE 2 demonstrates that the most pronounced negative shift in d15N from pre-OAE 2 to OAE 2 occurs in the open ocean, but with d15N never lower than -3 ppm. Using a box model of N cycling for the proto-North Atlantic during OAE 2, we show that N2-fixation is a major contributor to the d15N signal, especially in the open ocean. Incomplete uptake of ammonium for phytoplankton growth is important in regions dominated by downwelling, with lateral transport of ammonium acting as a major source. In the southern proto-North Atlantic, where bottom waters were euxinic, the light d15N signature is largely explained by upwelling of ammonium . Our study provides an overview of regional differences in d15N in the proto-North Atlantic and highlights the role of lateral exchange of water and nutrients, in addition to local biogeochemical processes, in determining d15N values of OAE 2 sediments.

Upwelling and isolation in oxygen-depleted anticyclonic modewater eddies and implications for nitrate cycling

The physical (temperature, salinity, velocity) and biogeochemical (oxygen, nitrate) structure of an oxygen depleted coherent, baroclinic, anticyclonic mode-water eddy (ACME) is investigated using high-resolution autonomous glider and ship data. A distinct core with a diameter of about 70 km is found in the eddy, extending from about 60 to 200 m depth and. The core is occupied by fresh and cold water with low oxygen and high nitrate concentrations, and bordered by local maxima in buoyancy frequency. Velocity and property gradient sections show vertical layering at the flanks and underneath the eddy characteristic for vertical propagation (to several hundred-meters depth) of near inertial internal waves (NIW) and confirmed by direct current measurements. A narrow region exists at the outer edge of the eddy where NIW can propagate downward. NIW phase speed and mean flow are of similar magnitude and critical layer formation is expected to occur. An asymmetry in the NIW pattern is seen that possible relates to the large-scale Ekman transport interacting with ACME dynamics. NIW/mean flow induced mixing occurs close to the euphotic zone/mixed layer and upward nutrient flux is expected and supported by the observations. Combing high resolution nitrate (NO3-) data with the apparent oxygen utilization (AOU) reveals AOU:NO3- ratios of 16 which are much higher than in the surrounding waters (8.1). A maximum NO3- deficit of 4 to 6 µmol kg-1 is estimated for the low oxygen core. Denitrification would be a possible explanation. This study provides evidence that the recycling of NO3-, extracted from the eddy core and replenished into the core via the particle export, may quantitatively be more important. In this case, the particulate phase is of keys importance in decoupling the nitrogen from the oxygen cycling.