A loss-of-function mutation of rice DENSE PANICLE 1 causes semi-dwarfness and slightly increased number of spikelets

In cereal breeding, semi-dwarfness and an increased spikelet number are favorable characteristics. We show that the rice DENSE PANICLE 1 (DN1) mutant allele Dn1-1 causes both of these characteristics and that Dn1-1 is a loss-of-function mutation. DN1 is allelic to DENSE AND ERECT PANICLE 1 (DEP1) (=qPE(9-1)). The expression level of OsCKX2 in the shoot apex of Dn1-1 plants is similar to that in the wild type, indicating that OsCKX2 does not contribute to an increased number of spikelets. A comparison of the Dn1-1 and Dn1-3 alleles suggests that the N-terminal region of DN1 contains a coiled-coil domain and a nuclear localization signal that might be responsible for semi-dwarfness. This comparison also revealed that a single transmembrane alpha-helix, a VWFC module, and a four-disulfide core domain can further increase spikelet number. Subcellular localization analysis of the DN1 protein fused with green fluorescent protein (GFP) implies that DN1 is located in the nucleus and cell membrane and that its N-terminal fragment is cleaved. Dn1-1 plants have normal sensitivity to gibberellin, brassinolide, and kinetin, and we observed no genetic epistasis with brassinolide-related mutants, suggesting that DN1 does not function in the signaling pathways of these phytohormones.