The maize SBP-box transcription factor encoded by tasselsheath4 regulates bract development and the establishment of meristem boundaries

Plant architecture consists of repeating units called phytomers, each containing an internode, leaf and axillary meristem. The formation of boundaries within the phytomer is necessary to differentiate and separate these three components, otherwise some will grow at the expense of others. The microRNA-targeted SBP-box transcription factor tasselsheath4 (tsh4) plays an essential role in establishing these boundaries within the inflorescence. tsh4 mutants display altered phyllotaxy, fewer lateral meristems and ectopic leaves that grow at the expense of the meristem. Double-mutant analyses of tsh4 and several highly branched mutants, such as ramosa1-3 and branched silkless1, demonstrated a requirement for tsh4 in branch meristem initiation and maintenance. TSH4 protein, however, was localized throughout the inflorescence stem and at the base of lateral meristems, but not within the meristem itself. Double labeling of TSH4 with the ramosa2, branched silkless1 and knotted1 meristem markers confirmed that TSH4 forms a boundary adjacent to all lateral meristems. Indeed, double labeling of miR156 showed a meristem-specific pattern complementary to that of TSH4, consistent with tsh4 being negatively regulated by this microRNA. Thus, downregulation of TSH4 by a combination of microRNAs and branching pathway genes allows the establishment of lateral meristems and the repression of leaf initiation, thereby playing a major role in defining meristem versus leaf boundaries.