Phylogenomic analyses of the BARREN STALK1/LAX PANICLE1 (BA1/LAX1) genes and evidence for their roles during axillary meristem development

The diversity of plant architectural form is largely determined by the extent and duration of axillary meristem (AM) derived lateral growth. The orthologous basic helix-loop-helix (bHLH) proteins maize BARREN STALK1 (BA1) and rice LAX PANICLE1 (LAX1) are essential for the formation of AMs during vegetative development and all lateral structures during inflorescence development, but whether BA1/LAX1 co-orthologs exist outside of the grass family is unclear. Here, we present Bayesian phylogenetic evidence of a well-supported BA1/LAX1 clade comprised monocots and eudicots, estimating an origin for the lineage at least near the base of flowering plants. Genomic analyses in Arabidopsis, papaya, medicago, rice, sorghum, and maize indicate that BA1/LAX1 genes reside in syntenic regions, although there has also been a complex pattern of gene duplication and loss during the diversification of the angiosperm clade. BA1/LAX1 mRNA expression coincided with the initiation of leaves and associated AMs in the vegetative meristems of broccoli, medicago, and papaya implicating a role for the lineage in the formation of AMs in eudicots as well as monocots. Expression on the adaxial surface of lateral inflorescence structures was conserved in all sampled flowering plants, whereas mRNA expression in leaves of Arabidopsis, broccoli, and papaya also links BA1/LAX1 co-orthologs with roles in regulating leaf development, possibly as a downstream target of auxin regulating genes. Together these data point to roles for BA1/LAX1 genes during AM formation, leaf, and inflorescence development in diverse flowering plants and lend support to the hypothesis that the same genetic mechanisms regulate the development of different AM types.