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Journal of Bacteriology, November 2001, p. 6454-6465, Vol. 183, No. 21
0021-9193/01/$04.00+0   DOI: 10.1128/JB.183.21.6454-6465.2001
Copyright © 2001, American Society for Microbiology. All rights reserved.

Functional Analysis of Genes for Biosynthesis of Pyocyanin and Phenazine-1-Carboxamide from Pseudomonas aeruginosa PAO1

Dmitri V. Mavrodi,1,2,* Robert F. Bonsall,1 Shannon M. Delaney,2 Marilyn J. Soule,2 Greg Phillips,3 and Linda S. Thomashow1,2,3

Department of Plant Pathology1 and USDA-ARS Root Disease and Biological Control Research Unit, Agricultural Research Service,3 Washington State University, Pullman, Washington 99164-6430, and School of Molecular Biosciences, Washington State University, Pullman, Washington 99164-42342

Received 27 April 2001/Accepted 18 July 2001

Two seven-gene phenazine biosynthetic loci were cloned from Pseudomonas aeruginosa PAO1. The operons, designated phzA1B1C1D1E1F1G1 and phzA2B2C2D2E2F2G2, are homologous to previously studied phenazine biosynthetic operons from Pseudomonas fluorescens and Pseudomonas aureofaciens. Functional studies of phenazine-nonproducing strains of fluorescent pseudomonads indicated that each of the biosynthetic operons from P. aeruginosa is sufficient for production of a single compound, phenazine-1-carboxylic acid (PCA). Subsequent conversion of PCA to pyocyanin is mediated in P. aeruginosa by two novel phenazine-modifying genes, phzM and phzS, which encode putative phenazine-specific methyltransferase and flavin-containing monooxygenase, respectively. Expression of phzS alone in Escherichia coli or in enzymes, pyocyanin-nonproducing P. fluorescens resulted in conversion of PCA to 1-hydroxyphenazine. P. aeruginosa with insertionally inactivated phzM or phzS developed pyocyanin-deficient phenotypes. A third phenazine-modifying gene, phzH, which has a homologue in Pseudomonas chlororaphis, also was identified and was shown to control synthesis of phenazine-1-carboxamide from PCA in P. aeruginosa PAO1. Our results suggest that there is a complex pyocyanin biosynthetic pathway in P. aeruginosa consisting of two core loci responsible for synthesis of PCA and three additional genes encoding unique enzymes involved in the conversion of PCA to pyocyanin, 1-hydroxyphenazine, and phenazine-1-carboxamide.

* Corresponding author. Mailing address: Department of Plant Pathology, P.O. Box 646430, Washington State University, Pullman, WA 99164-6430. Phone: (509) 335-3269. Fax: (509) 335-7674. E-mail: mavrodi{at}mail.wsu.edu.

Journal of Bacteriology, November 2001, p. 6454-6465, Vol. 183, No. 21
0021-9193/01/$04.00+0   DOI: 10.1128/JB.183.21.6454-6465.2001
Copyright © 2001, American Society for Microbiology. All rights reserved.


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