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. 2018 Aug 21;8:12504. doi: 10.1038/s41598-018-30884-x

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© The Author(s) 2018

Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/.

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Changes of fluxes for changing oxygen uptake by the symbiont. Grey values indicate no feasible solution (i.e. insufficient oxygen for sufficient nitrogen fixation). The upper row corresponds to wild type solutions while the lower row represents alanine dehydrogenase knockout. NH3 export is only active in the mutated strain, while all nitrogen is exported as alanine in the wild type simulation (marked in orange, as the export does not change with oxygen in both conditions). With increasing oxygen availability, succinate is preferentially used as carbon source as it provides additional reductant compared to Malate. Amino acid shuttling is reduced with increasing oxygen in both conditions, due to the increased potential of complete respiration of carbon. Under oxygen limiting conditions, pyruvate is generated instead of citrate, which can either be converted to alanine (wild type), consuming reductant, or can be used as acceptor for the amino group from glutamate.