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The outer edges of the Lzr gene cluster were defined based on comparisons to the BE-54017 and cladoniamide gene clusters and a BLAST analysis of genes surrounding the core indolotryptoline biosynthesis genes (Fig. 3A). The biosynthesis of indolotryptolines is well-character- ized, making it possible to predict the function of most genes in the Lzr gene cluster (32). The four key stages of indolotryptoline biosynthesis involve dimerization of oxo-tryptophan to form a chromopyrrolic acid, oxidative aryl–aryl coupling to form an indolocarbazole, “flipping” of one of the indole rings to form an indolotryptoline, and tailoring to generate the final product. The Lzr gene cluster is predicted to contain seven transcriptional units controlled by three bidirectional (P1, P2, and P3) and one unidirectional (P4) promoter regions (Fig. 3 B, i). This cluster is conveniently organized such that successive activation of the three bidirectional promoter regions (P1, P2, and P3) is predicted to drive the expression of genes required to achieve the first, second, and third stages in indolotryptoline biosynthesis, respectively (Fig. 3C). |
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