The plant growth hormone auxin plays a critical role in the initiation of lateral organs and meristems. Here, we identify and characterize a mutant, sparse inflorescence1 (spi1), which has defects in the initiation of axillary meristems and lateral organs during vegetative and inflorescence development in maize. Positional cloning shows that spi1 encodes a flavin monooxygenase similar to the YUCCA (YUC) genes of Arabidopsis, which are involved in local auxin biosynthesis in various plant tissues. In Arabidopsis, loss of function of single members of the YUC family has no obvious effect, but in maize the mutation of a single yuc locus causes severe developmental defects. Phylogenetic analysis of the different members of the YUC family in moss, monocot, and eudicot species shows that there have been independent expansions of the family in monocots and eudicots. spi1 belongs to a monocot-specific clade, within which the role of individual YUC genes has diversified. These observations, together with expression and functional data, suggest that spi1 has evolved a dominant role in auxin biosynthesis that is essential for normal maize inflorescence development. Analysis of the interaction between spi1 and genes regulating auxin transport indicate that auxin transport and biosynthesis function synergistically to regulate the formation of axillary meristems and lateral organs in maize.auxin biosynthesis ͉ auxin transport ͉ yucca ͉ meristem T he plant hormone auxin is required for initiation and polar growth of all organ primordia. Auxin is synthesized by a number of pathways in the cell and is transported from cell to cell by diffusion and by the activity of influx and efflux carriers (1). During vegetative development, auxin is required for many developmental processes, including leaf and lateral root initiation, whereas during inflorescence development, auxin is required for the initiation of floral meristems (FMs) and floral organs (2-5). Extensive research in Arabidopsis has shown the importance of auxin transport during lateral organ and axillary meristem initiation (3-5). In addition, recent work has highlighted the role of localized auxin biosynthesis in all aspects of plant development (6-10). The role of auxin in monocots such as maize is not as well understood. Although some aspects of the control of auxin transport seem to be conserved between monocots and eudicots (11-16), there are also key differences (17).Maize plants produce separate male and female inflorescences (18). The male inflorescence, the tassel, is situated at the shoot apex, whereas the female inflorescence, the ear, is produced from an axillary meristem several nodes below the tassel. The tassel consists of a main spike with several long lateral branches at the base ( Fig. 1 A and B). Both the main spike and branches are covered with short branches, each of which bears a pair of spikelets. Each spikelet produces two leaf-like glumes that enclose a pair of florets. Florets consist of a lemma and palea (outer whorl structures derived from bracts...

Auxin plays a fundamental role in organogenesis in plants. Multiple pathways for auxin biosynthesis have been proposed, but none of the predicted pathways are completely understood. Here, we report the positional cloning and characterization of the vanishing tassel2 (vt2) gene of maize (Zea mays). Phylogenetic analyses indicate that vt2 is a co-ortholog of TRYPTOPHAN AMINOTRANSFERASE OF ARABIDOPSIS1 (TAA1), which converts Trp to indole-3-pyruvic acid in one of four hypothesized Trp-dependent auxin biosynthesis pathways. Unlike single mutations in TAA1, which cause subtle morphological phenotypes in Arabidopsis thaliana, vt2 mutants have dramatic effects on vegetative and reproductive development. vt2 mutants share many similarities with sparse inflorescence1 (spi1) mutants in maize. spi1 is proposed to encode an enzyme in the tryptamine pathway for Trp-dependent auxin biosynthesis, although this biochemical activity has recently been questioned. Surprisingly, spi1 vt2 double mutants had only a slightly more severe phenotype than vt2 single mutants. Furthermore, both spi1 and vt2 single mutants exhibited a reduction in free auxin levels, but the spi1 vt2 double mutants did not have a further reduction compared with vt2 single mutants. Therefore, both spi1 and vt2 function in auxin biosynthesis in maize, possibly in the same pathway rather than independently as previously proposed.

Maize (Zea mays) has a highly branched inflorescence due to the production of different types of axillary meristems. Characterization of the barren inflorescence class of mutants has led to the discovery of genes required for axillary meristem initiation in the inflorescence. Previous studies showed that barren inflorescence2 (bif2) encodes a serine/threonine protein kinase that regulates auxin transport, and barren stalk1 (ba1) encodes a basic helix-loop-helix transcription factor that acts downstream of auxin transport. Here, we characterize Barren inflorescence1 (Bif1), a classical semidominant mutation of maize. Developmental, histological, and genetic analyses show that Bif1 mutants are defective in the initiation of all axillary meristems in the inflorescence. Real time RT-PCR experiments show that both bif2 and ba1 are expressed at lower levels in Bif1 mutants. Double-mutant analyses demonstrate that Bif1 exhibits an epistatic interaction with ba1 and a synergistic interaction with bif2. The dramatic phenotypic enhancement observed in Bif1; bif2 double mutants implies that bif1 plays an overlapping role with bif2 in the initiation of lateral organs during vegetative development. The phenotypic resemblance of Bif1 to bif2 mutants and the reduction of auxin transport in Bif1 mutants suggest that bif1 functions as a regulator of auxin transport in maize.

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The sparse inflorescence1 (spi1), Barren inflorescence1 (Bif1), barren inflorescence2 (bif2), and barren stalk1 (ba1) mutants produce fewer branches and spikelets in the inflorescence due to defects in auxin biosynthesis, transport, or response. We report that spi1, bif1, and ba1, but not bif2, also function in promoting cell elongation in the inflorescence.