Inflorescence architecture in small-grain cereals has a direct effect on yield and is an important selection target in breeding for yield improvement. We analyzed the recessive mutation laxatum-a (lax-a) in barley (Hordeum vulgare), which causes pleiotropic changes in spike development, resulting in (1) extended rachis internodes conferring a more relaxed inflorescence, (2) broadened base of the lemma awns, (3) thinner grains that are largely exposed due to reduced marginal growth of the palea and lemma, and (4) and homeotic conversion of lodicules into two stamenoid structures. Map-based cloning enforced by mapping-by-sequencing of the mutant lax-a locus enabled the identification of a homolog of BLADE-ON-PETIOLE1 (BOP1) and BOP2 as the causal gene. Interestingly, the recently identified barley uniculme4 gene also is a BOP1/2 homolog and has been shown to regulate tillering and leaf sheath development. While the Arabidopsis (Arabidopsis thaliana) BOP1 and BOP2 genes act redundantly, the barley genes contribute independent effects in specifying the developmental growth of vegetative and reproductive organs, respectively. Analysis of natural genetic diversity revealed strikingly different haplotype diversity for the two paralogous barley genes, likely affected by the respective genomic environments, since no indication for an active selection process was detected.The inflorescence is the most prominent part of smallgrain cereal plants, producing the carbohydrate-rich grains that are harvested for food, feed, and fiber. However, our understanding of the genetic factors that regulate inflorescence architecture remains limited. What is clear is that the appearance and shape of the inflorescence has been under constant visual selection since early domestication and is still ongoing in modern plant breeding due to the impact of inflorescence architecture on crop yield. For instance, in barley (Hordeum vulgare), strong selection has been exerted on spontaneously occurring alleles of nonbrittle rachis1 (btr1) and btr2 that prevent dehiscence of the rachis at maturity (Pourkheirandish et al., 2015), six-rowed spike1 (vrs1) that determines whether the inflorescence exhibits two or six rows of grain (Komatsuda et al., 2007), and nudum (nud) that controls whether the grain is hulled or hull-less (Taketa et al., 2008). Ultimately, knowing all of the genes that control cereal inflorescence architecture will provide targets for understanding and exploiting natural or induced genetic diversity toward improving both yield potential and end-use characteristics.The barley inflorescence or spike forms an unbranched main rachis carrying triplets of sessile single-floreted