The Rice G Protein γ Subunit DEP1/qPE9–1 Positively Regulates Grain-Filling Process by Increasing Auxin and Cytokinin Content in Rice Grains

Heterotrimeric G protein-mediated signal transduction is one of the most important and highly conserved signaling pathways in eukaryotes, which involves in the regulation of many important biological processes. As compared with those in mammals and Arabidopsis thaliana, the functions of rice heterotrimeric G protein and their molecular mechanisms are largely unknown. The rice genome contains a single G(α) (RGA1) and G(β) (RGB1), and five G(γ) (RGG1, RGG2, GS3, DEP1/qPE9–1, and GGC2) subunits. Recent genetic studies have shown that DEP1/qPE9–1, an atypical putative G(γ) protein, is responsible for the grain size as well as the dense and erect panicles, but the biochemical and molecular mechanisms underlying the control of grain size are not well understood. Here, we report that rice plants carrying DEP1/qPE9–1 have more endosperm cells per grain than plants contain the dep1/qpe9–1 allele. The DEP1/qPE9–1 line has a higher rate and more prolonged period of starch accumulation than the dep1/qpe9–1 line. Additionally, the expression of several essential genes encoding enzymes catalyzing sucrose metabolism and starch biosynthesis is higher in the DEP1/qPE9–1 line than in the dep1/qpe9–1 line, especially from the mid to late grain-filling stage. Grains of the DEP1/qPE9–1 line also have higher contents of three phytohormones, ABA, auxin and cytokinin. Exogenous application of auxin or cytokinin enhanced the starch accumulation and the expression of genes encoding grain-filling-related enzymes in the grains of dep1/qpe9–1, whereas ABA produced no effects. Based on these results, we conclude that DEP1/qPE9–1 positively regulates starch accumulation primarily through auxin and cytokinin, which enhance the expression of genes encoding starch biosynthesis during the mid to late grain-filling stage, resulting in increased duration of the grain-filling process.