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MBE Advance Access published online on December 23, 2003

Molecular Biology and Evolution, doi:10.1093/molbev/msh047
Molecular Biology and Evolution © Society for Molecular Biology and Evolution 2003; all rights reserved
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Accepted October 25, 2003
© 2003 Society for Molecular Biology and Evolution

Original Articles

The Natural History of Nitrogen Fixation

Jason Raymond 1, Janet L. Siefert 2, Christopher R. Staples 1, and Robert E. Blankenship 1*

1 Department of Chemistry and Biochemistry, Arizona State University, Tempe, AZ 85287-1604, USA
2 Department of Statistics, Rice University, Houston, TX 77251-1892, USA

* To whom correspondence should be addressed. E-mail: blankenship{at}asu.edu.


   Abstract

In recent years our understanding of biological nitrogen fixation has been bolstered by a diverse array of scientific techniques. Still, the origin and extant distribution of nitrogen fixation has been perplexing from a phylogenetic perspective, largely due to factors that confound molecular phylogeny such as sequence divergence, paralogy, and horizontal gene transfer. Here we make use of 110 publicly available complete genome sequences to understand how the core components of nitrogenase, including Nif H, D, K, E, and N proteins, have evolved. These genes are universal in nitrogen fixing organisms - typically found within highly conserved operons - and, overall, have remarkably congruent phylogenetic histories. Additional clues to the early origins of this system are available from two distinct clades of nitrogenase paralogs: one group comprised of genes essential to photosynthetic pigment biosynthesis, and an additional group of uncharacterized genes present in methanogens and some photosynthetic bacteria. We explore the complex genetic history of the nitrogenase family, which is replete with gene duplication, recruitment, fusion, and horizontal gene transfer, and discuss these events in light of the hypothesized presence of nitrogenase in the last common ancestor of modern organisms, as well as the additional possibility that nitrogen fixation might have evolved later, perhaps in methanogenic archaea, and was subsequently transferred into the bacterial domain.

Key Words: nitrogen fixation, nitrogenase, evolution, horizontal gene transfer


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