Increased planting densities have boosted maize yields. Upright plant architecture facilitates dense planting. Here, we cloned UPA1 (Upright Plant Architecture1) and UPA2, two quantitative trait loci conferring upright plant architecture. UPA2 is controlled by a two-base sequence polymorphism regulating the expression of a B3-domain transcription factor (ZmRAVL1) located 9.5 kilobases downstream. UPA2 exhibits differential binding by DRL1 (DROOPING LEAF1), and DRL1 physically interacts with LG1 (LIGULELESS1) and represses LG1 activation of ZmRAVL1. ZmRAVL1 regulates brd1 (brassinosteroid C-6 oxidase1), which underlies UPA1, altering endogenous brassinosteroid content and leaf angle. The UPA2 allele that reduces leaf angle originated from teosinte, the wild ancestor of maize, and has been lost during maize domestication. Introgressing the wild UPA2 allele into modern hybrids and editing ZmRAVL1 enhance high-density maize yields.