Combined Nitrogen

  • John A. RavenEmail author
  • Mario GiordanoEmail author
Part of the Developments in Applied Phycology book series (DAPH, volume 6)


Algae can use a wide range of combined N sources. All of them can use NH4+, and probably also use urea and one of more amino acids; most of them can also use NO2 and NO3, and some can use betaines and/or purines. Transport of combined N into cells very often uses H+ or Na+ symport. Two cations transported per anionic N species, and one cation transported per neutral N species, enables the electrical potential difference across the membrane generated by active cation efflux to be used to increase the accumulation ratio inside:outside of the combined N species. Cationic N forms, e.g. NH4+, sometimes occur at very low concentrations in the natural environment, and cation symport can increase the steady-state NH4+ concentration. The transporters have been to some extent characterised at the molecular level, especially for the plasmalemma. Assimilation of inorganic N species into organic N within the cell use well-established pathways, i.e. NO3 reductase, NO2 reductase, and glutamine synthetase – glutamine-oxoglutarate aminotransferase enzymes. N assimilation, and especially the initial step (NO3 reductase), are under more direct redox control in microalgae than in vascular plants. Combined N species which as NO and NO3 are involved in signalling within the cell, but extent to which they modulate metabolism in response to internal and external cues needs clarification. It is important to bear in mind that the conclusions drawn here generally come from work on relatively few microalgal species, and generalisations should be viewed with caution.


Amino acids Ammonium Betaines Combined nitrogen Organic nitrogen Nitrate Nitric oxide Nitrite Proton symport Purines Signalling Sodium symport Urea 



The University of Dundee is a registered Scottish charity, No SC 015096. MG’s research on N and S was funded by the Italian Ministry for Agriculture (MIPAF, Bioforme project), by the Italian Ministry of Foreign Affairs (MAE, Joint Italian-Israel Cooperation Program) and by the Assemble program of the European Union. Discussions with Mitchell Andrews, Hans Lambers and Bill Plaxton have been very helpful.


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© Springer International Publishing Switzerland 2016

Authors and Affiliations

  1. 1.Division of Plant BiologyUniversity of Dundee at the James Hutton InstituteInvergowrieUK
  2. 2.Plant Functional Biology and Climate Change ClusterUniversity of Technology SydneyUltimoAustralia
  3. 3.Laboratorio di Fisiologia delle Alghe e delle Piante, Dipartimento di Scienze della Vita e dell’AmbienteUniversità Politecnica delle MarcheAnconaItaly

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