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BIOLOGICAL SCIENCES / GENETICS
Enzymes of the shikimic acid pathway encoded in the genome of a basal metazoan, Nematostella vectensis, have microbial origins

Antonio Starcevic^*,, Shamima Akthar, Walter C. Dunlap, J. Malcolm Shick^¶, Daslav Hranueli^*, John Cullum, and Paul F. Long^,||

*Section for Bioinformatics, Department of Biochemical Engineering, Faculty of Food Technology and Biotechnology, University of Zagreb, Pierottijeva 6, 10000 Zagreb, Croatia; Department of Genetics, University of Kaiserslautern, Postfach 3049, 67653 Kaiserslautern, Germany; School of Pharmacy, University of London, 29/39 Brunswick Square, London WC1N 1AX, United Kingdom; Australian Institute of Marine Science, PMB 3 Townsville MC, Townsville 4810, Queensland, Australia; and ^¶School of Marine Sciences, University of Maine, 5751 Murray Hall, Orono, ME 04469-5751

Edited by Lynn Margulis, University of Massachusetts, Amherst, MA, and approved December 19, 2007 (received for review August 7, 2007)

The shikimic acid pathway is responsible for the biosynthesis of many aromatic compounds by a broad range of organisms, including bacteria, fungi, plants, and some protozoans. Animals are considered to lack this pathway, as evinced by their dietary requirement for shikimate-derived aromatic amino acids. We challenge the universality of this traditional view in this report of genes encoding enzymes for the shikimate pathway in an animal, the starlet sea anemone Nematostella vectensis. Molecular evidence establishes horizontal transfer of ancestral genes of the shikimic acid pathway into the N. vectensis genome from both bacterial and eukaryotic (dinoflagellate) donors. Bioinformatic analysis also reveals four genes that are closely related to those of Tenacibaculum sp. MED152, raising speculation for the existence of a previously unsuspected bacterial symbiont. Indeed, the genome of the holobiont (i.e., the entity consisting of the host and its symbionts) comprises a high content of Tenacibaculum-like gene orthologs, including a 16S rRNA sequence that establishes the phylogenetic position of this associate to be within the family Flavobacteriaceae. These results provide a complementary view for the biogenesis of shikimate-related metabolites in marine Cnidaria as a "shared metabolic adaptation" between the partners.

symbiosis | Tenacibaculum | Cnidaria

The authors declare no conflict of interest.

This article is a PNAS Direct Submission.

This article contains supporting information online at www.pnas.org/cgi/content/full/0707388105/DC1.

^||To whom correspondence should be addressed. E-mail: paul.long{at}pharmacy.ac.uk

© 2008 by The National Academy of Sciences of the USA

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