Axillary branch suppression is a favorable trait bred into many domesticated crop plants including maize compared with its highly branched wild ancestor teosinte. Branch suppression in maize was achieved through selection of a gain of function allele of the teosinte branched1 (tb1) transcription factor that acts as a repressor of axillary bud growth. Previous work indicated that other loci may function epistatically with tb1 and may be responsible for some of its phenotypic effects. Here, we show that tb1 mediates axillary branch suppression through direct activation of the tassels replace upper ears1 (tru1) gene that encodes an ankyrin repeat domain protein containing a BTB/POZ motif necessary for protein-protein interactions. The expression of TRU1 and TB1 overlap in axillary buds, and TB1 binds to two locations in the tru1 gene as shown by chromatin immunoprecipitation and gel shifts. In addition, nucleotide diversity surveys indicate that tru1, like tb1, was a target of selection. In modern maize, TRU1 is highly expressed in the leaf trace vasculature of axillary internodes, while in teosinte, this expression is highly reduced or absent. This increase in TRU1 expression levels in modern maize is supported by comparisons of relative protein levels with teosinte as well as by quantitative measurements of mRNA levels. Hence, a major innovation in creating ideal maize plant architecture originated from ectopic overexpression of tru1 in axillary branches, a critical step in mediating the effects of domestication by tb1.A xillary meristems are produced at the base of each new leaf and can have different fates depending on their positions within the plant. In maize, those found aboveground have a reproductive fate, while those found at the base of the plant can become long branches that are vegetative clones of the main shoot and are called tillers. These extra branches often compete for valuable resources such as light (1) and can have a negative impact on plant growth if produced in excess. Indeed, suppression of axillary branching through an increase in apical dominance is a commonly bred trait among many domesticated crop plants (2), including maize (Zea mays ssp. mays), which was domesticated from teosinte (Z. mays ssp. parviglumis), its wild ancestor. While there are multiple genetic pathways necessary for suppression of axillary bud production in plants (3-5), in maize, this process has been found to operate through a specialized pathway mediated by the teosinte branched1 (tb1) locus first identified by Charles Burnham over 50 y ago (6). tb1 loss of function mutants overproduce tillers and have long aerial branches tipped by male tassels that replace the normally female ears (7), indicating that it functions as a repressor of both axillary bud growth and inflorescence sexual fate (8). Previous quantitative trait locus studies mapping the genetic basis for differences in plant morphology between maize and teosinte showed that selection at the tb1 locus was responsible for the increase in apical dominance foun...