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The maize PI gene encodes a myb-homologous protein that regulates flavonoid pigment biosynthesis in floral organs including the kernel pericarp, cob glume and tassel glume. The Pl-rr allele coding sequence is flanked by long (5.2 kbp) direct repeats. To clarily the origin of the unusual P-rr gene structure, we isolated and characterized a second, tightly linked, maize gene {P2). and a related gene {P2-t) from teosinte parviglumis. the closest wild relative of maize. The maize P2 gene and the teosinte P2-t gene are homologous to the maize PI gene in the coding region and 3" flanking region, but not in the 5' flanking region. The 5" flanking regions of maize P2 and teosinte P2-t are similar to each other, but different from that of the maize PI gene. We propose that the maize PI and P2 genes were generated by a tandem (head-to-tail) gene duplication in which the 3" flanking sequence of the progenitor P gene was duplicated at the 5' side of the incipient PI gene, thereby generating the long direct repeats that flank the Pl-rr allele. This duplication event occurred approximately 2.75 MYA. and subsequent insertions of retroelements separated the PI and P2 genes. Pigmentation phenotypes and transcript analysis indicate that the two duplicated maize P genes and the teosinte P2-t gene have distinct tissue specific expression patterns. The

Molecular characterization of myb-homologous transcriptional factors of the flavonoid pathway in Zea mays