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Nitrogen fixation in the Arctic Ocean

Lead Research Organisation: University of Liverpool
Department Name: Earth, Ocean and Ecological Sciences

Abstract

Biological nitrogen fixation is the conversion of nitrogen gas (N2) to fixed nitrogen (e.g. nitrate). N2 fixation is a crucial component of global ocean biogeochemical cycles. It provides the major source of nitrogen necessary to balance nitrogen loss via denitrification and annamox and thus controls the magnitude of fixed nitrogen in the ocean, with consequences for the cycling and storage of carbon and other nutrients. Until recently, N2 fixation was thought to be restricted to the warm surface waters (20 to 30dC) of the low latitude subtropical gyres. This perception has focused decades of research on nitrogen fixation to the low latitude ocean, where other nutrients such as iron and phosphorus limit the activity of nitrogen fixers.

However, there is new evidence that nitrogen fixing organisms are present and active in the cold surface waters of the Arctic Ocean. The nitrogen fixing gene has been detected in the western and central Arctic Ocean and observed rates of nitrogen are comparable to warm water nitrogen fixation. This unexpected result means that our current understanding of global nitrogen fixation is incomplete as it neglects to account for nitrogen fixation in cold waters. In addition, we do not know how temperature, light, nutrients and iron control the distribution and activity of nitrogen fixing organisms in the Arctic Ocean. There is an urgent need to better understand nitrogen fixation in the Arctic Ocean for three reasons. Firstly, the primary productivity of the Arctic Ocean is already limited by the availability of nitrate. Warming of the Arctic Ocean has caused sea ice to decline by 9% per decade since the 1970s, causing primary productivity to increase by 25%. As the Arctic Ocean warms and becomes ice free, primary production is predicted to increase. To support increased primary production, there must be an additional source of nitrate, but this source remains elusive. Inputs of nitrate from rivers, the atmosphere and dissolved organic nitrogen do not meet the nitrate demand associated with increased primary production. Nitrogen fixation may therefore be a crucial source of nitrogen in the contemporary and future Arctic Ocean and provide the fixed nitrogen necessary to support the future increase in primary production. Secondly, there is ongoing debate on the status of the nitrogen budget in the Arctic Ocean. Although the amount of nitrate entering and leaving the Arctic Ocean is equal, there is a large nitrate sink in the sediments on the Arctic shelves. This implies that either there is a large deficit in nitrate in the Arctic Ocean, or there is currently a source of nitrate that is unaccounted for. Nitrogen fixation may be the missing source required to balance the Arctic Ocean nitrogen budget. However, the lack of observations means that pan-Arctic estimates of nitrogen fixation are rudimentary. Finally, global numerical models that represent the oceanic nutrient cycling and the marine ecosystem currently ignore nitrogen fixation in the Arctic Ocean. This means that the ability of these models to predict how the Arctic ocean will respond to increased warming and increased productivity is inaccurate because they don't account for the additional nutrient source or the effect nitrogen fixation has on nutrients such as carbon, phosphorus and iron. Overall, our lack of knowledge on nitrogen fixation in the Arctic Ocean means we do not know how important this process is to global nitrogen fixation or nitrogen budget.

N-ARC proposes to conduct the first holistic study of nitrogen fixation in the eastern Arctic Ocean. We chose this region for two reasons; (a) new data from N-ARC team has detected the gene responsible for nitrogen fixation in this region, providing new evidence that nitrogen fixation is occurring in the eastern Arctic Ocean and (b) there are strong gradients in temperature and nutrients, allowing us to explore how the ocean conditions control nitrogen fixation in the Arctic Ocean.

Planned Impact

Who might benefit from this research?

The main beneficiaries of this project will be science leaders and stakeholders such as the Arctic Council concerned with climate change in the Arctic, school pupils, teachers and wider general public.

How might they benefit from this research?
Science leaders and policy makers: The Arctic Ocean is receiving an incredible amount of international attention due to the dramatic and rapid environmental changes observed over the past few decades, and the uncertainty in the response of the Arctic and its ecosystem as the environment continues to warm. Despite the wave of research activity across the Arctic, there are still key processes, such as nitrogen fixation, not being quantified or missing from climate models used to predict the future direction and magnitude of change in productivity and thus ecosystem health in the Arctic. N-ARC will tackle a fundamental problem in the Arctic ocean, that is, to assess a potentially overlooked source of nutrients that may be critical to sustain primary production now and in the future. Our observational findings will provide new insight into how nitrogen fixation shapes contemporary and future Arctic Ocean nutrient cycles and productivity. Our outputs will thus provide a basis to improve predictions on how the future Arctic ocean may respond to a changing climate. We believe these findings will be of interest not only to Arctic scientists, but policy makers interested in climate change in the Arctic. N-ARC directly addresses NERC strategy to 'Understand and predict how our planet works'.

School children and the general public; It is essential that our research outcomes are disseminated to the UK community. This is challenging but vital if we are to recruit students into sciences, convince the public that it is worthwhile funding this type of research and increase awareness of the sensitivity of the Arctic to a changing climate. We propose to communicate science from N-ARC in two ways. Firstly, there are key topics within the school curriculum for GCSEs and A-levels that relate directly to N-ARC, such as the nitrogen cycle. Our project will develop 'tool boxes' containing curriculum relevant material to demonstrate key concepts in our research project and these will be made available to school teachers. In addition, we have experience in engaging with school children and the general public via events such as Pint of Science, Big Bang NW, Meet the Scientist and Mersey River Festival and will continue to use these events to communicate with a large audience.

We will work with the University of Liverpool Marketing and Communications team to create a series of short (2 minute) videos describing (a) climate change in the Arctic Ocean, (b) the problem of the missing nutrient source and (c) findings from N-ARC. The videos would be prepared from the start of the project, thus capturing Arctic fieldwork and would be released alongside the first high profile paper.

We will hire two graduates from a science communication course at the University of Liverpool for 3 months to develop a toolbox to disseminate the motivation and findings from N-ARC to school children based on national curriculum and to STEM events as above.

Publications

10 25 50
 
Description We found evidence for nitrogen fixation in the Arctic Ocean, specifically rates of nitrogen fixation similar to those found in the warm subtropical ocean, and molecular evidence for the existence of nitrogen fixing microbes in the Arctic Ocean. We discovered higher nitrogen fixation associated with glacier outflow and sea ice, and also with particles.
We conducted nutrient addition experiments to investigate which nutrient limits phytoplankton growth in the Arctic Ocean and found evidence of widespread nitrate limitation, alongside evidence of co-limitation with phosphorus and iron.
Exploitation Route The data will be added to global databases on nitrogen fixation (rates and genes) and also help develop functional relationships to improve representation of nitrogen fixation in global models.
Sectors Environment

 
Title DY167 Calculations for nitrogen fixation rates in the North Sea including limits of detection (LOD) and minimum quantifiable rates (MQR) 
Description Calculations of rates of nitrogen fixation from samples collected during DY167 to the North Sea during July 2023 onboard RRS Discovery. Calculations are based on those published by Gradoville et al 2017 and include estimates of the limits of detection and minimum quantifiable rates. 
Type Of Material Database/Collection of data 
Year Produced 2025 
Provided To Others? Yes  
Impact NA 
URL https://datacat.liverpool.ac.uk/id/eprint/3033
 
Title DY167 North Sea Surfmet data including nutrients, iron, manganese and particulate carbon 
Description The dataset consists of measurements taken from the surface waters (< 3m) in the North Sea in July 2023. The measurements include sea surface temperature (°C), sea surface salinity, chlorophyll a concentrations (µg per L) and transmission (%) from calibrated sensors. Discrete water samples were collected using a trace metal clean towed FISH. Filtered seawater samples were analysed for inorganic nutrients including nitrate+nitrite, phosphate and silicate (µM), dissolved iron (nM) and dissolved manganese (nM). Samples were filtered for the analysis of particulate carbon concentrations (µM). 
Type Of Material Database/Collection of data 
Year Produced 2025 
Provided To Others? Yes  
Impact NA 
URL https://datacat.liverpool.ac.uk/id/eprint/3039
 
Title Nitrate+nitrite, phosphate, silicate, chlorophyll a, Fv:Fm data from nutrient addition incubation experiments during cruise DY167 in the Barents Sea, July-August 2023 
Description Nutrient addition experiments were performed during research cruise DY167 to the Barents Sea between 9th July and 13th August 2023. Six experiments were conducted, the first as a test station (T1) then five at locations representing different water characteristics, specifically Atlantic water (N03), ice edge (N08), glacier influenced (N15) and polar waters north of the polar front (N07 and N07Y). Twenty four 20L carboys were filled with unfiltered seawater using seawater collected from the 12th of July to the 6th of August using the trace metal clean towed-FISH. Nitrate, phosphate and iron chloride were added alone or in combination to a final concentration of 10 µM, 0.8 µM and 2 nM. Experimental setup and subsampling was performed under trace metal clean conditions. Experiments were performed in triplicate (3 carboys per control or treatment). Carboys were incubated in a temperature and light controlled refrigerated container, with incubation temperature set to sea surface temperature. LED light panels (Part no: LED-PANEL-300-1200-DW and LED-PANEL-200-6-DW, Daylight White, supplier Power Pax UK Limited) delivered a photon flux ranging from 180 to 220 µmol photons m-2 s-1 and were on 24 hours to mimic light at this time of year in the Arctic. Carboys were subsampled for nutrients, chlorophyll a, and variable fluorescence vs maximum fluorescence (Fv:Fm) at time zero and then once a day for the duration of the experiment. Experiments were conducted for between 4 and 7 days depending on the rate of change of chlorophyll and Fv:Fm. Nutrients were measured using standard colorimetric methods and a Seal Analytical QuAAtro 5 channel autoanalyser. Chlorophyll a was measured using acetone extraction and fluorimetric analysis via a Turner 10Au fluorometer. Fv:Fm values were calculated from measurements made using a Chelsea Technology Group FRRF II Sensor. The experiments were conducted to investigate the macro and micronutrient control on primary productivity in the Barents Sea, under Natural Environment Research Council (NERC) Discovery grant Nitrogen fixation in the Arctic Ocean, N-ARC (grant numbers NE/T000570/1 and NE/T001240/1). 
Type Of Material Database/Collection of data 
Year Produced 2025 
Provided To Others? Yes  
Impact NA 
URL https://www.bodc.ac.uk/data/published_data_library/catalogue/10.5285/3bda959d-9b1f-7486-e063-7086abc...
 
Title Raw ddPCR abundance of UCYN-A2 and UCYN-A1 from DY167 
Description Abundances of B. bigelowii/UCYN-A1 and B. bigelowii//UCYN-A2/A3/A4 were estimated from 11 surface water filtered seawater samples (< 3m, Sterivex 0.22 µm)using digital droplet PCR (ddPCR) using the assays originally developed for Taqman® quantitative PCR by Church et al., (2005) and Thompson et al., (2014), respectively. All reaction conditions, thermocycling parameters, thresholding and calculations of the limit of detection and limit of quantitation are described in Gradoville et al., (2020). Note that the abundances determined using the UCYN-A2/A3/A4 assay likely represent both UCYN-A2 and UCYN-A4, both of which were recovered via nifH amplicon HTS. 
Type Of Material Database/Collection of data 
Year Produced 2025 
Provided To Others? Yes  
Impact NA 
URL https://datacat.liverpool.ac.uk/id/eprint/3040
 
Title Stainless steel CTD water column profiles (temperature, salinity, dissolved oxygen, optical backscatter, beam transmittance and attenuation, PAR, chlorophyll-a fluorescence ) collected in the Norwegian and Barents Sea in July-August 2023 on cruise... 
Description This dataset contains in-situ temperature, salinity, dissolved oxygen, optical backscatter, beam transmittance and attenuation, PAR, chlorophyll-a fluorescence data from 186 downcast stainless steel CTD profiles. The measurements were collected during cruise DY167 (GEOTRACES cruise GApr19) in July-August 2023 in the Norwegian and Barents Seas and cover the relatively warm Atlantic Water inflow, fresher water on the Spitzbergen Bank, coastal/glacial outflow around Svalbard, and Arctic water around sea ice. The stainless steel unit comprised a Sea-Bird Electronics (SBE) 9 plus underwater unit, and an SBE 11 plus deck unit; all of which were mounted on a stainless steel 24-way carousel. Attached to the underwater unit were two SBE 3P temperature sensors, two SBE 4C conductivity sensors, one Paroscientific Digiquartz pressure sensor, two SBE 43 dissolved oxygen sensors, two Biospherical QCP Cosine PAR sensors, one WETLabs BBRTD light scattering sensor, one Valeport altimeter one CTG Aquatracka MKIII fluorometer and WETLabs C-star transmissometer. Raw data were initially extracted and processed using Seabird Data Processing Software. Modules DatCnv, WildEdit, Filter, AlignCTD, CellTM and Derive were all run. Custom Matlab routines were then used to remove out of water and surface soak values, to filter anomalous data points and to average the profiles onto a 1 decibar pressure grid. Salinity, chlorophyll-a, and oxygen concentrations were all calibrated against independent in-situ samples. These data were collected to better determine how temperature, light, nutrients, and iron control the distribution and activity of nitrogen fixing organisms in the Arctic Ocean. This dataset was generated by the National Oceanography Centre (NOC) under Natural Environment Research Council (NERC) Discovery grant Nitrogen fixation in the Arctic Ocean, N-ARC (grant numbers NE/T000570/1 and NE/T001240/1). 
Type Of Material Database/Collection of data 
Year Produced 2026 
Provided To Others? Yes  
Impact NA 
URL https://www.bodc.ac.uk/data/published_data_library/catalogue/10.5285/4c1edb44-81e5-b3ad-e063-7086abc...
 
Title Trace metal free CTD water column profiles (temperature, salinity, dissolved oxygen, optical backscatter, beam transmittance and attenuation, PAR, pH, and chlorophyll-a fluorescence) collected in the Norwegian and Barents Sea in July-August 2023 on... 
Description This dataset contains in-situ temperature, salinity, dissolved oxygen, optical backscatter, beam transmittance and attenuation, PAR, pH, and chlorophyll-a fluorescence data from 22 downcast trace-metal free CTD profiles. The measurements were collected during cruise DY167 (GEOTRACES cruise GApr19) in July-August 2023 in the Norwegian and Barents Seas and cover the relatively warm Atlantic Water inflow, fresher water on the Spitzbergen Bank, coastal/glacial outflow around Svalbard, and Arctic water around sea ice. The trace metal free setup comprised a Sea-Bird Electronics (SBE) 9 plus underwater unit, and an SBE 11 plus deck unit; all of which were mounted on a titanium 24-way carousel. Attached to the underwater unit were two SBE 3P temperature sensors, two SBE 4C conductivity sensors, one Paroscientific Digiquartz pressure sensor, one SBE 43 dissolved oxygen sensor, two CTG PAR sensors, one WETLabs BBRTD light scattering sensor, one Valeport altimeter, one CTG Aquatracka MKIII fluorometer, one WETLabs C-star transmissometer and one AMT Deep pH sensor. Raw data were initially extracted and processed using Seabird Data Processing Software. Modules DatCnv, WildEdit, Filter, AlignCTD, CellTM and Derive were all run. Custom Matlab routines were then used to remove out of water and surface soak values, to filter anomalous data points and to average the profiles onto a 1 decibar pressure grid. Salinity and chlorophyll-a were both calibrated against independent in-situ samples. These data were collected to better determine how temperature, light, nutrients, and iron control the distribution and activity of nitrogen fixing organisms in the Arctic Ocean. This dataset was generated by the National Oceanography Centre (NOC) under Natural Environment Research Council (NERC) Discovery grant Nitrogen fixation in the Arctic Ocean, N-ARC (grant numbers NE/T000570/1 and NE/T001240/1). 
Type Of Material Database/Collection of data 
Year Produced 2026 
Provided To Others? Yes  
Impact NA 
URL https://www.bodc.ac.uk/data/published_data_library/catalogue/10.5285/4c1d29b9-ae76-6619-e063-7086abc...
 
Description Collaboration with Mar Benavides from the Mediterranean Institute of Oceanography 
Organisation Mediterranean Institute of Oceanography
Country France 
Sector Public 
PI Contribution Invitation to participate in a research cruise to the Arctic Ocean (DY167) in summer 2023. Provision of consumables to support research on the ship. Access to ancillary data sets (physical oceanography, nutrients etc).
Collaborator Contribution Participation in research cruise with PhD student Arthur Coet Measurement of particle associated nitrogen fixation and diazotroph community structure
Impact Particle-Associated Diazotrophs in the Changing Arctic Ocean, Poster Presentation at 2024 Ocean Sciences Meeting USA
Start Year 2022
 
Description Collaboration with Prof. Nicolas Cassar, Duke University USA 
Organisation Duke University
Country United States 
Sector Academic/University 
PI Contribution Prof. Nicolas Cassar is a project partner on the grant and has received financial contribution from Liverpool of £150,000
Collaborator Contribution Use of state of the art instruments on the research cruise to measure nitrogen fixation in high resolution and in near real time.
Impact Data sets from Duke University have been used in a large grant outline to NERC
Start Year 2023