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  • Review Article
  • Published:

Emerging patterns of marine nitrogen fixation

Nature Reviews Microbiology volume 9, pages 499–508 (2011)Cite this article

Subjects

Key Points

  • N2 fixation in the ocean is an important process that contributes to the biological sequestration of CO2 in the deep ocean.

  • There are different types of marine diazotrophs (including heterotrophic bacteria), but those that are known to contribute to marine N2 fixation are: members of the genus Trichodesmium, which are non-heterocystous, filamentous cyanobacteria; the unicellular cyanobacteria UCYN-A and Crocosphaera watsonii; free-living heterocystous species such as Nodularia spp. and Anabaena spp.; and the heterocystous diatom symbiont Richelia intracellularis. Although all of these are cyanobacteria, it is predicted that N2 fixation in UCYN-A is dependent on organic C (that is, UCYN-A is a secondary producer), whereas the others are primary producers.

  • The temperature defines the boundaries of where each type of organism can be found, and the deposition of dust (and in the case of the Baltic Sea the runoff from land), the nutrient supply and the internal cycling of nutrients control which nutrients (Fe or P) will be limiting to diazotrophs. Diazotrophs have developed a number of physiological adaptations to deal with both types of nutrient limitation.

  • Fe and P also seem to control the distribution of diazotrophic species in the major ocean basins, with the high Fe areas of the North Atlantic Ocean and Arabian Sea hosting high abundances of Trichodesmium spp., whereas the North and South Pacific Ocean and South Atlantic Ocean seem to be dominated by unicellular cyanobacterial diazotrophs. The distribution of diazotrophic species is important, and different morphological and physiological types have the potential to affect the mode of transfer of newly fixed N into the food web and the sequestration of atmospheric CO2 in the deep sea.

  • Research is now focusing on how increased atmospheric CO2 affects N2 fixation, and on the search for N2 fixation in waters where it has not been previously considered: that is, lower-temperature waters and waters with measureable amounts of fixed inorganic N. High CO2 can increase N2 fixation in Trichodesmium spp. and C. watsonii.

Abstract

Biological N2 fixation is an important part of the marine nitrogen cycle as it provides a source of new nitrogen that can support biological carbon export and sequestration. Research in the past decade has focused on determining the patterns of distribution and abundance of diazotrophs, defining the environmental features leading to these patterns and characterizing the factors that constrain marine N2 fixation overall. In this Review, we describe how variations in the deposition of iron from dust to different ocean basins affects the limiting nutrient for N2 fixation and the distribution of different diazotrophic species. However, many questions remain about marine N2 fixation, including the role of temperature, fixed nitrogen species, CO2 and physical forcing in controlling N2 fixation, as well as the potential for heterotrophic N2 fixation.

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Figure 1: Overview of the importance of nitrogen fixation in the ocean.
Figure 2: Summary of the distribution of nitrogen fixation rates, diazotrophic species and nutrient limitation in the ocean.

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Acknowledgements

The authors would like to express their thanks to M. Church, R. Foster, R. Langlois, J. LaRoche, P. Moisander and J. Zehr for providing Trichodesmium spp. nifH data and J. K. Moore for a model output, both of which were included in a previous version of this manuscript. Three anonymous reviewers helped to greatly improve the manuscript.

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  1. Department of Biological Sciences and Wrigley Institute for Environmental Studies, University of Southern California, Los Angeles, 90089, California, USA

    Jill A. Sohm, Eric A. Webb & Douglas G. Capone

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  1. Jill A. Sohm
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Glossary

Upwelling

Wind-mediated movement of deep water to the surface.

Physical forcing

The effect of physical conditions and processes in the ocean on biological properties.

Diazotrophs

Organisms that can fix N2 by converting it to ammonia.

Heterocyst

A specialized cell with a thick cell wall that lacks photosystem II (PSII) and is the site of N2 fixation in some filamentous cyanobacteria.

Blooms

Areas of large growth or accumulation of a species.

Brackish

Describing water that has a salinity between fresh and marine water.

Dissolved Fe

Fe that can pass through a 0.4 μm filter.

Free Fe

A pool of Fe that is composed of Fe bound to inorganic ligands and a small amount of free ion (Fe3+) and is presumed to be bioavailable (often denoted Fe′).

Ligand

In this article, an organic molecule that binds Fe (potentially a siderophore).

Sulpholipid

A membrane lipid containing a sulphur (as opposed to P)-based head.

Alkaline phosphatase

A hydrolytic enzyme that cleaves phosphomonoesters from P-containing dissolved organic matter.

Blackman limitation

The limitation of the growth rate of an organism.

Liebig limitation

The limitation of the yield of an organism (as in a crop yield).

Gyre

A giant circular surface current that is present in the ocean.

Luxury uptake

The uptake of nutrients that are in excess of demand, and that can be made into storage products.

Eddies

Small-scale circular currents in which the water inside is typically different from the water outside.

C concentrating mechanism

The mechanism that increases the concentration of CO2 around ribulose-1,5-bisphosphate carboxylase oxygenase (RuBisCo), in order to increase the rate of photosynthesis.

Anticyclonic eddies

Eddies with warm interior water and a centre that is slightly higher than the surrounding sea surface. They rotate clockwise in the northern hemisphere and anticlockwise in the southern hemisphere.

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Sohm, J., Webb, E. & Capone, D. Emerging patterns of marine nitrogen fixation. Nat Rev Microbiol 9, 499–508 (2011). https://doi.org/10.1038/nrmicro2594

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