scholarly journals Dynamics of N<sub>2</sub> fixation and fate of diazotroph-derived nitrogen in a low nutrient low chlorophyll ecosystem: results from the VAHINE mesocosm experiment (New Caledonia)

2015 ◽  
Vol 12 (23) ◽  
pp. 19579-19626 ◽  
Author(s):  
S. Bonnet ◽  
H. Berthelot ◽  
K. Turk-Kubo ◽  
S. Fawcett ◽  
E. Rahav ◽  
...  

Abstract. N2 fixation rates were measured daily in large (~ 50 m3) mesocosms deployed in the tropical South West Pacific coastal ocean (New Caledonia) to investigate the spatial and temporal dynamics of diazotrophy and the fate of diazotroph-derived nitrogen (DDN) in a low nutrient, low chlorophyll ecosystem. The mesocosms were intentionally fertilized with ~ 0.8 μM dissolved inorganic phosphorus (DIP) to stimulate diazotrophy. Bulk N2 fixation rates were replicable between the three mesocosms, averaged 18.5 ± 1.1 nmol N L−1 d−1 over the 23 days, and increased by a factor of two during the second half of the experiment (days 15 to 23) to reach 27.3 ± 1.0 nmol N L−1 d−1. These rates are higher than the upper range reported for the global ocean, indicating that the waters surrounding New Caledonia are particularly favourable for N2 fixation. During the 23 days of the experiment, N2 fixation rates were positively correlated with seawater temperature, primary production, bacterial production, standing stocks of particulate organic carbon, nitrogen and phosphorus, and alkaline phosphatase activity, and negatively correlated with DIP concentrations, DIP turnover time, nitrate, and dissolved organic nitrogen and phosphorus concentrations. The fate of DDN was investigated during the bloom of the unicellular diazotroph, UCYN-C, that occurred during the second half of the experiment. Quantification of diazotrophs in the sediment traps indicates that ~ 10 % of UCYN-C from the water column were exported daily to the traps, representing as much as 22.4 ± 5.5 % of the total POC exported at the height of the UCYN-C bloom. This export was mainly due to the aggregation of small (5.7 ± 0.8 μm) UCYN-C cells into large (100–500 μm) aggregates. During the same time period, a DDN transfer experiment based on high-resolution nanometer scale secondary ion mass spectrometry (nanoSIMS) coupled with 15N2 isotopic labelling revealed that 16 ± 6 % of the DDN was released to the dissolved pool and 21 ± 4 % was transferred to non-diazotrophic plankton, mainly picoplankton (18 ± 4 %) followed by diatoms (3 ± 2 %) within 24 h of incubation. This is consistent with the observed dramatic increase in picoplankton and diatom abundances, primary production, bacterial production and standing stocks of particulate organic carbon, nitrogen and phosphorus during the second half of the experiment in the mesocosms. These results offer insights into the fate of DDN during a bloom of UCYN-C in low nutrient, low chlorophyll ecosystems.

2016 ◽  
Vol 13 (9) ◽  
pp. 2653-2673 ◽  
Author(s):  
Sophie Bonnet ◽  
Hugo Berthelot ◽  
Kendra Turk-Kubo ◽  
Sarah Fawcett ◽  
Eyal Rahav ◽  
...  

Abstract. N2 fixation rates were measured daily in large (∼ 50 m3) mesocosms deployed in the tropical southwest Pacific coastal ocean (New Caledonia) to investigate the temporal variability in N2 fixation rates in relation with environmental parameters and study the fate of diazotroph-derived nitrogen (DDN) in a low-nutrient, low-chlorophyll ecosystem. The mesocosms were fertilized with  ∼ 0.8 µM dissolved inorganic phosphorus (DIP) to stimulate diazotrophy. Bulk N2 fixation rates were replicable between the three mesocosms, averaged 18.5 ± 1.1 nmol N L−1 d−1 over the 23 days, and increased by a factor of 2 during the second half of the experiment (days 15 to 23) to reach 27.3 ± 1.0 nmol N L−1 d−1. These later rates measured after the DIP fertilization are higher than the upper range reported for the global ocean. During the 23 days of the experiment, N2 fixation rates were positively correlated with seawater temperature, primary production, bacterial production, standing stocks of particulate organic carbon (POC), nitrogen (PON) and phosphorus (POP), and alkaline phosphatase activity, and negatively correlated with DIP concentrations, DIP turnover time, nitrate, and dissolved organic nitrogen and phosphorus concentrations. The fate of DDN was investigated during a bloom of the unicellular diazotroph UCYN-C that occurred during the second half of the experiment. Quantification of diazotrophs in the sediment traps indicates that ∼ 10 % of UCYN-C from the water column was exported daily to the traps, representing as much as 22.4 ± 5.5 % of the total POC exported at the height of the UCYN-C bloom. This export was mainly due to the aggregation of small (5.7 ± 0.8 µm) UCYN-C cells into large (100–500 µm) aggregates. During the same time period, a DDN transfer experiment based on high-resolution nanometer-scale secondary ion mass spectrometry (nanoSIMS) coupled with 15N2 isotopic labeling revealed that 16 ± 6 % of the DDN was released to the dissolved pool and 21 ± 4 % was transferred to non-diazotrophic plankton, mainly picoplankton (18 ± 4 %) followed by diatoms (3 ± 2 %). This is consistent with the observed dramatic increase in picoplankton and diatom abundances, primary production, bacterial production, and standing stocks of POC, PON, and POP in the mesocosms during the second half of the experiment. These results offer insights into the fate of DDN during a bloom of UCYN-C in low-nutrient, low-chlorophyll ecosystems.


2016 ◽  
Vol 13 (21) ◽  
pp. 6081-6093 ◽  
Author(s):  
Kristian Spilling ◽  
Kai G. Schulz ◽  
Allanah J. Paul ◽  
Tim Boxhammer ◽  
Eric P. Achterberg ◽  
...  

Abstract. About a quarter of anthropogenic CO2 emissions are currently taken up by the oceans, decreasing seawater pH. We performed a mesocosm experiment in the Baltic Sea in order to investigate the consequences of increasing CO2 levels on pelagic carbon fluxes. A gradient of different CO2 scenarios, ranging from ambient ( ∼  370 µatm) to high ( ∼  1200 µatm), were set up in mesocosm bags ( ∼  55 m3). We determined standing stocks and temporal changes of total particulate carbon (TPC), dissolved organic carbon (DOC), dissolved inorganic carbon (DIC), and particulate organic carbon (POC) of specific plankton groups. We also measured carbon flux via CO2 exchange with the atmosphere and sedimentation (export), and biological rate measurements of primary production, bacterial production, and total respiration. The experiment lasted for 44 days and was divided into three different phases (I: t0–t16; II: t17–t30; III: t31–t43). Pools of TPC, DOC, and DIC were approximately 420, 7200, and 25 200 mmol C m−2 at the start of the experiment, and the initial CO2 additions increased the DIC pool by  ∼  7 % in the highest CO2 treatment. Overall, there was a decrease in TPC and increase of DOC over the course of the experiment. The decrease in TPC was lower, and increase in DOC higher, in treatments with added CO2. During phase I the estimated gross primary production (GPP) was  ∼  100 mmol C m−2 day−1, from which 75–95 % was respired,  ∼  1 % ended up in the TPC (including export), and 5–25 % was added to the DOC pool. During phase II, the respiration loss increased to  ∼  100 % of GPP at the ambient CO2 concentration, whereas respiration was lower (85–95 % of GPP) in the highest CO2 treatment. Bacterial production was  ∼  30 % lower, on average, at the highest CO2 concentration than in the controls during phases II and III. This resulted in a higher accumulation of DOC and lower reduction in the TPC pool in the elevated CO2 treatments at the end of phase II extending throughout phase III. The “extra” organic carbon at high CO2 remained fixed in an increasing biomass of small-sized plankton and in the DOC pool, and did not transfer into large, sinking aggregates. Our results revealed a clear effect of increasing CO2 on the carbon budget and mineralization, in particular under nutrient limited conditions. Lower carbon loss processes (respiration and bacterial remineralization) at elevated CO2 levels resulted in higher TPC and DOC pools than ambient CO2 concentration. These results highlight the importance of addressing not only net changes in carbon standing stocks but also carbon fluxes and budgets to better disentangle the effects of ocean acidification.


2015 ◽  
Vol 12 (23) ◽  
pp. 19861-19900 ◽  
Author(s):  
F. Van Wambeke ◽  
U. Pfreundt ◽  
A. Barani ◽  
H. Berthelot ◽  
T. Moutin ◽  
...  

Abstract. N2 fixation fuels ~ 50 % of new primary production in the oligotrophic South Pacific Ocean. The VAHINE mesocosm experiment designed to track the fate of diazotroph derived nitrogen (DDN) in the New Caledonia lagoon. Here, we examined the temporal dynamics of heterotrophic bacterial production during this experiment. Three replicate large-volume (~ 50 m3) mesocosms were deployed and were intentionally fertilized with dissolved inorganic phosphorus (DIP) to stimulate N2 fixation. We specifically examined relationships between N2 fixation rates and primary production, determined bacterial growth efficiency and established carbon budgets of the system from the DIP fertilization to the end of the experiment (days 5–23). Heterotrophic bacterioplankton production (BP) and alkaline phosphatase activity (APA) were statistically higher during the second phase of the experiment (P2: days 15–23), when chlorophyll biomass started to increase compared to the first phase (P1: days 5–14). Among autotrophs, Synechococcus abundances increased during P2, possibly related to its capacity to assimilate leucine and to produce alkaline phosphatase. Bacterial growth efficiency based on the carbon budget was notably higher than generally cited for oligotrophic environments (27–43 %), possibly due to a high representation of proteorhodopsin-containing organisms within the picoplanctonic community. The carbon budget showed that the main fate of gross primary production (particulate + dissolved) was respiration (67 %), and export through sedimentation (17 %). BP was highly correlated with particulate primary production and chlorophyll biomass during both phases of the experiment but slightly correlated, and only during P2 phase, with N2 fixation rates. Our results suggest that most of the DDN reached the heterotrophic bacterial community through indirect processes, like mortality, lysis and grazing.


2016 ◽  
Author(s):  
Kristian Spilling ◽  
Kai G. Schulz ◽  
Allanah J. Paul ◽  
Tim Boxhammer ◽  
Eric P. Achterberg ◽  
...  

Abstract. About a quarter of anthropogenic CO2 emissions are currently taken up by the oceans decreasing seawater pH. We performed a mesocosm experiment in the Baltic Sea in order to investigate the consequences of increasing CO2 levels on pelagic carbon fluxes. A gradient of different CO2 scenarios, ranging from ambient (~ 370 µatm) to high (~ 1200 µatm), were set up in mesocosm bags (~ 55 m3). We determined standing stocks and temporal changes of total particulate carbon (TPC), dissolved organic (DOC), dissolved inorganic (DIC) and particulate organic carbon (POC) of specific plankton groups. We also measured carbon flux via CO2 exchange with the atmosphere and sedimentation (export); and biological rate measurements of primary production, bacterial production and total respiration. The experiment lasted for 44 days and was divided into three different phases (I: t0–t16; II: t17–t30; III: t31–t43). Pools of TPC, DOC and DIC were approximately 420, 7200 and 25 200 mmol C m−2 at the start of the experiment, and the initial CO2 additions increased the DIC pool by ~ 7 % in the highest CO2 treatment. Overall, there was a decrease in TPC and increase of DOC over the course of the experiment. The decrease in TPC was lower, and increase in DOC higher, in treatments with added CO2. During Phase I the estimated gross primary production (GPP) was ~ 100 mmol C fixed m−2 d−1; from which 75–95 % were respired, ~ 1 % ended up in the TPC (including export) and 5–25 % added to the DOC pool. During Phase II, the respiration loss increased to ~ 100 % of GPP at the ambient CO2 concentration, whereas respiration was lower (85–95 % of GPP) in the highest CO2 treatment. Bacterial production was ~ 30 % lower, on average, at the highest CO2 concentration compared with the controls during Phases II and III. This resulted in a higher accumulation DOC standing stock and lower reduction in TPC in the elevated CO2 treatments at the end of Phase II extending throughout Phase III. The "extra" organic carbon at high CO2 remained fixed in an increasing biomass of small-sized plankton and in the DOC pool, and did not transferred into large, sinking aggregates. Our results revealed a clear effect of increasing CO2 on carbon production and mineralization, in particular under nutrient limited conditions. Lower carbon loss processes (respiration and bacterial remineralization) at elevated CO2 levels resulted in higher TPC and DOC pools compared with the ambient CO2 concentration. These results highlight the importance to address not only net changes in carbon standing stocks, but also carbon fluxes and budgets to better disentangle the effects of ocean acidification.


2016 ◽  
Vol 13 (11) ◽  
pp. 3187-3202 ◽  
Author(s):  
France Van Wambeke ◽  
Ulrike Pfreundt ◽  
Aude Barani ◽  
Hugo Berthelot ◽  
Thierry Moutin ◽  
...  

Abstract. Studies investigating the fate of diazotrophs through the microbial food web are lacking, although N2 fixation can fuel up to 50 % of new production in some oligotrophic oceans. In particular, the role played by heterotrophic prokaryotes in this transfer is largely unknown. In the frame of the VAHINE (VAriability of vertical and tropHIc transfer of diazotroph derived N in the south wEst Pacific) experiment, three replicate large-volume (∼ 50 m3) mesocosms were deployed for 23 days in the new Caledonia lagoon and were intentionally fertilized on day 4 with dissolved inorganic phosphorus (DIP) to stimulate N2 fixation. We specifically examined relationships between heterotrophic bacterial production (BP) and N2 fixation or primary production, determined bacterial growth efficiency and established carbon budgets. BP was statistically higher during the second phase of the experiment (P2: days 15–23), when chlorophyll biomass started to increase compared to the first phase (P1: days 5–14). Phosphatase alkaline activity increased drastically during the second phase of the experiment, showing adaptations of microbial populations after utilization of the added DIP. Notably, among autotrophs, Synechococcus abundances increased during P2, possibly related to its capacity to assimilate leucine and to produce alkaline phosphatase. Bacterial growth efficiency based on the carbon budget (27–43 %), was notably higher than generally cited for oligotrophic environments and discussed in links with the presence of abundant species of bacteria expressing proteorhodopsin. The main fates of gross primary production (particulate + dissolved) were respiration (67 %) and export through sedimentation (17 %). BP was highly correlated with particulate primary production and chlorophyll biomass during both phases of the experiment but was slightly correlated, and only during P2 phase, with N2 fixation rates. Heterotrophic bacterial production was strongly stimulated after mineral N enrichment experiments, suggesting N-limitation of heterotrophic bacteria across the experiment. N2 fixation rates corresponded to 17–37 % of the nitrogen demand of heterotrophic bacteria. Our results suggest that most of the diazotroph-derived nitrogen fuelled the heterotrophic bacterial community through indirect processes generating dissolved organic matter and detritus, like mortality, lysis and grazing of both diazotrophs and non-diazotrophs.


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