scholarly journals Nitrogen isotopic evidence for a shift from nitrate- to diazotroph-fueled export production in the VAHINE mesocosm experiments

2016 ◽  
Vol 13 (16) ◽  
pp. 4645-4657 ◽  
Author(s):  
Angela N. Knapp ◽  
Sarah E. Fawcett ◽  
Alfredo Martínez-Garcia ◽  
Nathalie Leblond ◽  
Thierry Moutin ◽  
...  

Abstract. In a coastal lagoon with a shallow, 25 m water column off the southwest coast of New Caledonia, large-volume ( ∼  50 m3) mesocosm experiments were undertaken to track the fate of newly fixed nitrogen (N). The mesocosms were intentionally fertilized with 0.8 µM dissolved inorganic phosphorus to stimulate diazotrophy. N isotopic evidence indicates that the dominant source of N fueling export production shifted from subsurface nitrate (NO3−) assimilated prior to the start of the 23-day experiments to N2 fixation by the end of the experiments. While the δ15N of the sinking particulate N (PNsink) flux changed during the experiments, the δ15N of the suspended PN (PNsusp) and dissolved organic N (DON) pools did not. This is consistent with previous observations that the δ15N of surface ocean N pools is less responsive than that of PNsink to changes in the dominant source of new N to surface waters. In spite of the absence of detectable NO3− in the mesocosms, the δ15N of PNsink indicated that NO3− continued to fuel a significant fraction of export production (20 to 60 %) throughout the 23-day experiments, with N2 fixation dominating export after about 2 weeks. The low rates of organic N export during the first 14 days were largely supported by NO3−, and phytoplankton abundance data suggest that sinking material primarily comprised large diatoms. Concurrent molecular and taxonomic studies indicate that the diazotroph community was dominated by diatom–diazotroph assemblages (DDAs) at this time. However, these DDAs represented a minor fraction (< 5 %) of the total diatom community and contributed very little new N via N2 fixation; they were thus not important for driving export production, either directly or indirectly. The unicellular cyanobacterial diazotroph, a Cyanothece-like UCYN-C, proliferated during the last phase of the experiments when N2 fixation, primary production, and the flux of PNsink increased significantly, and δ15N budgets reflected a predominantly diazotrophic source of N fueling export. At this time, the export flux itself was likely dominated by the non-diazotrophic diatom, Cylindrotheca closterium, along with lesser contributions from other eukaryotic phytoplankton and aggregated UCYN-C cells, as well as fecal pellets from zooplankton. Despite comprising a small fraction of the total biomass, UCYN-C was largely responsible for driving export production during the last  ∼  10 days of the experiments both directly ( ∼  5 to 22 % of PNsink) and through the rapid transfer of its newly fixed N to other phytoplankton; we infer that this newly fixed N was transferred rapidly through the dissolved N (including DON) and PNsusp pools. This inference reconciles previous observations of invariant oligotrophic surface ocean DON concentrations and δ15N with incubation studies showing that diazotrophs can release a significant fraction of their newly fixed N as some form of DON.

2015 ◽  
Vol 12 (23) ◽  
pp. 19901-19939 ◽  
Author(s):  
A. N. Knapp ◽  
S. E. Fawcett ◽  
A. Martínez-Garcia ◽  
N. Leblond ◽  
T. Moutin ◽  
...  

Abstract. In a shallow, coastal lagoon off the southwest coast of New Caledonia, large-volume (~ 50 m3) mesocosm experiments were undertaken to track the fate of newly fixed nitrogen (N). The mesocosms were intentionally fertilized with 0.8 μM dissolved inorganic phosphorus (DIP) to stimulate diazotrophy. N isotopic evidence indicates that the dominant source of N fueling export production shifted from subsurface nitrate (NO3−) assimilated prior to the start of the 23 day experiments to N2 fixation by the end of the experiments. While the δ15N of the sinking particulate N (PNsink) flux changed during the experiments, the δ15N of the suspended PN (PNsusp) and dissolved organic N (DON) pools did not. This is consistent with previous observations that the δ15N of surface ocean N pools is less responsive than that of PNsink to changes in the dominant source of new N to surface waters. In spite of the absence of detectable NO3− in the mesocosms, the δ15N of PNsink indicated that NO3− continued to fuel a significant fraction of export production (20 to 60 %) throughout the 23 day experiments, with N2 fixation dominating export after about two weeks. The low rates of primary productivity and export production during the first 14 days were primarily supported by NO3−, and phytoplankton abundance data suggest that export was driven by large diatoms sinking out of surface waters. Concurrent molecular and taxonomic studies indicate that the diazotroph community was dominated by diatom-diazotroph assemblages (DDAs) at this time. However, these DDAs represented a minor fraction (< 5 %) of the total diatom community and contributed very little new N via N2 fixation; they were thus not important for driving export production, either directly or indirectly. The unicellular cyanobacterial diazotroph, a Cyanothece-like UCYN-C, proliferated during the last phase of the experiments when N2 fixation, primary production, and the flux of PNsink increased significantly, and δ15N budgets reflected a predominantly diazotrophic source of N fueling export production. At this time, the export flux itself was likely dominated by the non-diazotrophic diatom, Cylindrotheca closterium, along with a lesser contribution from other eukaryotic phytoplankton and a small contribution (< 10 %) from aggregated UCYN-C cells. Despite comprising a small fraction of the total biomass, UCYN-C was largely responsible for driving export production during the last ~ 10 days of the experiments through the rapid transfer of its newly fixed N to other phytoplankton; we infer that this newly fixed N was transferred through the DON and/or ammonium pools. This inference reconciles previous observations of invariant oligotrophic surface ocean DON concentrations and δ15N with incubation studies showing that diazotrophs can release a significant fraction of their newly fixed N as some form of DON.


Author(s):  
Richard E. Zeebe ◽  
Andy Ridgwell

Over the period from 1750 to 2000, the oceans have absorbed about one-third of the carbon dioxide (CO2) emitted by humans. As the CO2 dissolves in seawater, the oceans become more acidic and between 1750 and 2000, anthropogenic CO2 emissions have led to a decrease of surface-ocean total pH (pH T) by ~0.1 units from ~8.2 to ~8.1 (see Chapters 1 and 3). Surface-ocean pHT has probably not been below ~8.1 during the past 2 million years (Hönisch et al. 2009). If CO2 emissions continue unabated, surface-ocean pH T could decline by about 0.7 units by 2300 (Zeebe et al. 2008). With increasing CO2 and decreasing pH, carbonate ion (CO32–) concentrations decrease and those of bicarbonate (HCO-3) rise. With declining CO32– concentration ([CO32–]), the stability of the calcium carbonate (CaCO3) mineral structure, used extensively by marine organisms to build shells and skeletons, is reduced. Other geochemical consequences include changes in trace metal speciation (Millero et al. 2009 ) and even sound absorption ( Hester et al. 2008 ; Ilyina et al. 2010 ). Do marine organisms and ecosystems really ‘care’ about these chemical changes? We know from a large number of laboratory, shipboard, and mesocosm experiments, that many marine organisms react in some way to changes in their geochemical environment like those that might occur by the end of this century (see Chapters 6 and 7). Generally (but not always), calcifying organisms produce less CaCO3, while some may put on more biomass. Extrapolating such experiments would lead us to expect potentially significant changes in ecosystem structure and nutrient cycling. But can one really extrapolate an instantaneous environmental change to one occurring on a timescale of a century? What capability, if any, do organisms have to adapt to future ocean acidification which is occurring on a slower timescale than can be replicated in the laboratory? Simultaneous changes in ocean temperature and nutrient supply as well as in organisms’ predation environment may create further stresses or work to ameliorate the effect of changes in ocean chemistry.


2016 ◽  
Vol 13 (13) ◽  
pp. 3901-3913 ◽  
Author(s):  
Allanah J. Paul ◽  
Eric P. Achterberg ◽  
Lennart T. Bach ◽  
Tim Boxhammer ◽  
Jan Czerny ◽  
...  

Abstract. Nitrogen fixation by filamentous cyanobacteria supplies significant amounts of new nitrogen (N) to the Baltic Sea. This balances N loss processes such as denitrification and anammox, and forms an important N source supporting primary and secondary production in N-limited post-spring bloom plankton communities. Laboratory studies suggest that filamentous diazotrophic cyanobacteria growth and N2-fixation rates are sensitive to ocean acidification, with potential implications for new N supply to the Baltic Sea. In this study, our aim was to assess the effect of ocean acidification on diazotroph growth and activity as well as the contribution of diazotrophically fixed N to N supply in a natural plankton assemblage. We enclosed a natural plankton community in a summer season in the Baltic Sea near the entrance to the Gulf of Finland in six large-scale mesocosms (volume ∼ 55 m3) and manipulated fCO2 over a range relevant for projected ocean acidification by the end of this century (average treatment fCO2: 365–1231 µatm). The direct response of diazotroph growth and activity was followed in the mesocosms over a 47 day study period during N-limited growth in the summer plankton community. Diazotrophic filamentous cyanobacteria abundance throughout the study period and N2-fixation rates (determined only until day 21 due to subsequent use of contaminated commercial 15N-N2 gas stocks) remained low. Thus estimated new N inputs from diazotrophy were too low to relieve N limitation and stimulate a summer phytoplankton bloom. Instead, regeneration of organic N sources likely sustained growth in the plankton community. We could not detect significant CO2-related differences in neither inorganic nor organic N pool sizes, or particulate matter N : P stoichiometry. Additionally, no significant effect of elevated CO2 on diazotroph activity was observed. Therefore, ocean acidification had no observable impact on N cycling or biogeochemistry in this N-limited, post-spring bloom plankton assemblage in the Baltic Sea.


2006 ◽  
Vol 18 (2) ◽  
pp. 333-339 ◽  
Author(s):  
André Luís Thomas ◽  
Ladaslav Sodek

The transport of organic N compounds to the shoot in the xylem sap of nodulated soybean plants was investigated in an attempt to better understand the changes in N metabolism under root hypoxia (first 5 days of flooding), with different sources of N in the medium. NO3- is beneficial for tolerance of plants to waterlogging, whereas other N sources such as NH4+ and NH4NO3, are not. Nevertheless, in the presence of NH4+ high levels of amino acids were transported in the xylem, consistent with its assimilation. Some increase in the transport of amino acids was also seen with NO3- nutrition during waterlogging, but not with N-free medium. Ureide transport in the xylem was severely reduced during waterlogging, consistent with impaired N2 fixation under these conditions. The relative proportions of some amino acids in the xylem showed dramatic changes during treatment. Alanine increased tremendously under root hypoxia, especially with NH4+ as N source, where it reached near 70 % of the total amino acids present. Aspartic acid, on the other hand, dropped to very low levels and was inversely related to alanine levels, consistent with this amino acid being the immediate source of N for alanine synthesis. Glutamine levels also fell to a larger or lesser extent, depending on the N source present. The changes in asparagine, one of the prominent amino acids of the xylem sap, were most outstanding in the treatment with NO3-, where they showed a large increase, characteristic of plants switching from dependence on N2 fixation to NO3- assimilation. The data indicate that the lesser effectiveness of NH4+ during waterlogging, in contrast to NO3-, involves restricted amino acids metabolism, and may result from energy metabolism being directed towards NH4+ detoxification.


2021 ◽  
pp. 117-141
Author(s):  
Jonathan P. Zehr ◽  
Douglas G. Capone
Keyword(s):  

1997 ◽  
Vol 24 (5) ◽  
pp. 631 ◽  
Author(s):  
O. Brendel ◽  
C. Wheeler ◽  
L. Handley

In field studies, 15N-enriched and 15N-natural abundance methods may yield similar mean estimates for N2-fixation, but with no correlation of the individual estimates. This study was designed as a glasshouse-based microcosm and aimed to remove the landform and weather variables found in previous field studies, while retaining other sources of variability such as mineralisation of organic N and whatever slight variations of environment may exist in the glasshouse. The results showed little or no correspondence among the estimates of N2-fixation at the three 15N-enrichment levels; however, estimates were consistent within each 15N-enrichment level. There was also no correlation between the results of acetylene reduction and those of any of the isotopic enrichment treatments. If the isotopic 15N-enrichment levels measured the same processes with varying accuracies, then the general patterns of the replicated results from each enrichment level should resemble each other against a generally uniform background. They did not. Hence, we conclude that the observed differences are due to unknown factors in addition to the varying noise-to-signal ratio at different enrichment levels.


2007 ◽  
Vol 4 (1) ◽  
pp. 75-86 ◽  
Author(s):  
M. A. Altabet

Abstract. According to current best estimates, the modern ocean's N cycle is in severe deficit. N isotope budgeting provides an independent geochemical constraint in this regard as well as the only means for past reconstruction. Overall, it is the relative proportion of N2 fixation consumed by water column denitrification that sets average oceanic δ15N under steady-state conditions. Several factors (conversion of organic N to N2, Rayleigh closed and open system effects) likely reduce the effective fractionation factor (ε) for water column denitrification to about half the inherent microbial value for εden. If so, the average oceanic δ15N of ~5‰ is consistent with a canonical contribution from water column denitrification of 50% of the source flux from N2 fixation. If an imbalance in oceanic N sources and sinks changes this proportion then a transient in average oceanic δ15N would occur. Using a simple model, changing water column denitrification by ±30% or N2 fixation by ±15% produces detectable (>1‰) changes in average oceanic δ15N over one residence time period or more with corresponding changes in oceanic N inventory. Changing sedimentary denitrification produces no change in δ15N but does change N inventory. Sediment δ15N records from sites thought to be sensitive to oceanic average δ15N all show no detectible change over the last 3 kyr or so implying a balanced marine N budget over the latest Holocene. A mismatch in time scales is the most likely meaningful interpretation of the apparent conflict with modern flux estimates. Decadal to centennial scale oscillations between net N deficit and net surplus may occur but on the N residence timescale of several thousand years, net balance is achieved in sum. However, sediment δ15N records from the literature covering the period since the last glacial maximum show excursions of up to several ‰ that are consistent with sustained N deficit during the deglaciation followed by readjustment and establishment of balance in the early Holocene. Since imbalance was sustained for one N residence time period or longer, excursions in ocean N inventory of 10 to 30% likely occurred. The climatic and oceanographic changes that occurred over this period evidently overcame, for a time, the capacity of ocean biogeochemistry to maintain N balance.


2018 ◽  
pp. 437-449
Author(s):  
Emese Szabó ◽  
Jakab Loch

The use of new methods describing the “readily available” nutrient content of the soil is spreading on a global scale. The 0.01 M CaCl2 extractant is a dilute salt solution in which the easily soluble inorganic (nitrate-N and ammonium-N) and organic N fractions, P, K and micronutrients are also measurable. The 0.01 M CaCl2 has been tested in the University of Debrecen, Institute of Agricultural Chemistry and Soil Sciences since the 90’s. The results of the researches related to organic N fraction, performed in the last decades, and the results of the present study (originating from the long-term experiment of Karcag, 2007–2009) can be concluded as follows: The measurement of easily soluble and oxidizable organic nitrogen (Norg), besides inorganic fractions, could improve the nutrient management. The amount of the Norg fraction is determined by the soil conditions, therefore it is considered to be a site-specific parameter. Management practices and cropyear affect the amount of Norg as well. The present research confirmed that, the effect of fertilization on the amount of Norg can be explained by the changing of the yield (related to total biomass production), while the effect of cropyear is related to the differences in mineralization circumstances and yield as well. The measurement of the Norg fraction is increases the accuracy of N-supply, therefore it could prevent the environmentally harmful excess N application as well.


2020 ◽  
Vol 13 (10) ◽  
pp. 4663-4690
Author(s):  
Markus Pahlow ◽  
Chia-Te Chien ◽  
Lionel A. Arteaga ◽  
Andreas Oschlies

Abstract. Uncertainties in projections of marine biogeochemistry from Earth system models (ESMs) are associated to a large degree with the imperfect representation of the marine plankton ecosystem, in particular the physiology of primary and secondary producers. Here, we describe the implementation of an optimality-based plankton–ecosystem model (OPEM) version 1.1 with variable carbon : nitrogen : phosphorus (C:N:P) stoichiometry in the University of Victoria ESM (UVic; Eby et al., 2009; Weaver et al., 2001) and the behaviour of two calibrated reference configurations, which differ in the assumed temperature dependence of diazotrophs. Predicted tracer distributions of oxygen and dissolved inorganic nutrients are similar to those of an earlier fixed-stoichiometry formulation in UVic (Nickelsen et al., 2015). Compared to the classic fixed-stoichiometry UVic model, OPEM is closer to recent satellite-based estimates of net community production (NCP), despite overestimating net primary production (NPP), can better reproduce deep-ocean gradients in the NO3-:PO43- ratio and partially explains observed patterns of particulate C:N:P in the surface ocean. Allowing diazotrophs to grow (but not necessarily fix N2) at similar temperatures as other phytoplankton results in a better representation of surface Chl and NPP in the Arctic and Antarctic oceans. Deficiencies of our calibrated OPEM configurations may serve as a magnifying glass for shortcomings in global biogeochemical models and hence guide future model development. The overestimation of NPP at low latitudes indicates the need for improved representations of temperature effects on biotic processes, as well as phytoplankton community composition, which may be represented by locally varying parameters based on suitable trade-offs. The similarity in the overestimation of NPP and surface autotrophic particulate organic carbon (POC) could indicate deficiencies in the representation of top-down control or nutrient supply to the surface ocean. Discrepancies between observed and predicted vertical gradients in particulate C:N:P ratios suggest the need to include preferential P remineralisation, which could also benefit the representation of N2 fixation. While OPEM yields a much improved distribution of surface N* (NO3--16⋅PO43-+2.9 mmol m−3), it still fails to reproduce observed N* in the Arctic, possibly related to a misrepresentation of the phytoplankton community there and the lack of benthic denitrification in the model. Coexisting ordinary and diazotrophic phytoplankton can exert strong control on N* in our simulations, which questions the interpretation of N* as reflecting the balance of N2 fixation and denitrification.


2007 ◽  
Vol 4 (1) ◽  
pp. 417-454 ◽  
Author(s):  
T. M. Lenton ◽  
C. A. Klausmeier

Abstract. We re-examine what controls the deep ocean N:P ratio in the light of recent findings that the C:N:P stoichiometry of phytoplankton varies with growth rate, nutrient and light limitation, species and phylum, and that N2-fixation may be limited by Fe or light in large parts of the world ocean. In particular, we assess whether a systematic change in phytoplankton stoichiometry can alter the deep ocean N:P ratio. To do this we adapt recent models to include non-Redfieldian stoichiometry of phytoplankton and restriction of N2-fixers to a fraction of the surface ocean. We show that a systematic change in phytoplankton C:N:P can alter the concentrations of NO3 and PO4 in the deep ocean but cannot greatly alter their ratio, unless it also alters the N:P threshold for N2-fixation. This occurs if competitive dynamics set the N:P threshold for N2-fixation, in which case it remains close to the N:P requirement of non-fixers (rather than that of N2-fixers) and consequently so does the deep ocean N:P ratio. Then, even if N2-fixers are restricted to a fraction of the surface ocean, they reach higher densities there, minimising variations in deep ocean N:P. Theoretical limits on the N:P requirements of phytoplankton suggest that since the deep ocean became well oxygenated, its N:P ratio is unlikely to have varied by more than a factor of two in either direction. Within these bounds, evolutionary changes in phytoplankton composition, and increased phosphorus weathering due to the biological colonisation of the land surface, are predicted to have driven long-term changes in ocean composition.


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