Within the cereal grasses, variation in inflorescence architecture results in a conspicuous morphological diversity that in crop species influences the yield of cereal grains. Although significant progress has been made in identifying some of the genes underlying this variation in maize and rice, in the temperate cereals, a group that includes wheat, barley, and rye, only the dosagedependent and highly pleiotropic Q locus in hexaploid wheat has been molecularly characterized. Here we show that the characteristic variation in the density of grains along the inflorescence, or spike, of modern cultivated barley (Hordeum vulgare) is largely the consequence of a perturbed interaction between microRNA172 and its corresponding binding site in the mRNA of an APELATA2 (AP2)-like transcription factor, HvAP2. We used genome-wide association and biparental mapping to identify HvAP2. By comparing inflorescence development and HvAP2 transcript abundance in an extreme dense-spike mutant and its nearly isogenic WT line, we show that HvAP2 turnover driven by microRNA 172 regulates the length of a critical developmental window that is required for elongation of the inflorescence internodes. Our data indicate that this heterochronic change, an altered timing of developmental events caused by specific temporal variation in the efficiency of HvAP2 turnover, leads to the striking differences in the size and shape of the barley spike.genome wide association scan | heterochrony T he cereal grasses are the world's most economically, sociologically, and ecologically important crops (1). Within and between species, variation in inflorescence architecture results in striking and characteristic morphological diversity. Dissecting the mechanisms and processes underpinning this diversity is central to understanding both its origin and evolution and how it can be harnessed for knowledge-based crop improvement. In maize and rice, a considerable body of work has begun to dissect the genes and gene networks controlling aspects of inflorescence development (e.g., refs. 2-5). However, within the Triticeae cereals, a group that includes some of the world's most important human foods, only Q, a highly pleiotropic gene that influences a broad range of phenotypic characters including inflorescence compactness, fragility, and free-threshing (6), and genes that determine the development of an extra floret on the lemma (7), additional glumes (8), the number of grains on the inflorescence (9), and floret fertility (10) have been molecularly characterized.Barley is a diploid inbreeding crop species and a model for other, genetically more complex temperate cereal species such as durum and bread wheats and rye. The barley inflorescence forms a terminal spike that bears reproductive units called "spikelets." Three spikelets initiate at each alternating node of the spike's central stem or rachis. Each spikelet forms a single floret that can develop into a kernel of grain. Several loci have been shown to affect spike density in barley (11), including dense spike...