Low phosphorus promotes NSP1-NSP2 heterodimerization to enhance strigolactone biosynthesis and regulate shoot and root architecture in rice.

Phosphorus is an essential macronutrient for plant development and metabolism, and plants have evolved ingenious mechanisms to overcome phosphate (Pi) starvation. However, the molecular mechanisms underlying the regulation of shoot and root architecture by low phosphorus conditions and the coordinated utilization of Pi and nitrogen remain largely unclear. Here, we show that Nodulation Signaling Pathway 1 (NSP1) and NSP2 regulate rice tiller number by promoting the biosynthesis of strigolactones (SLs), a class of phytohormones with fundamental effects on plant architecture and environmental responses. We found that NSP1 and NSP2 are induced by Oryza sativa PHOSPHATE STARVATION RESPONSE2 (OsPHR2) in response to low-Pi stress and form a complex to directly bind the promoters of SL biosynthesis genes, thus markedly increasing SL biosynthesis in rice. Interestingly, the NSP1/2-SL signaling module represses the expression of CROWN ROOTLESS 1 (CRL1), a newly identified early SL-responsive gene in roots, to restrain lateral root density under Pi deficiency. We also demonstrated that GR244DO treatment under normal conditions inhibits the expression of OsNRTs and OsAMTs to suppress nitrogen absorption but enhances the expression of OsPTs to promote Pi absorption, thus facilitating the balance between nitrogen and phosphorus uptake in rice. Importantly, we found that NSP1p:NSP1 and NSP2p:NSP2 transgenic plants show improved agronomic traits and grain yield under low- and medium-phosphorus conditions. Taken together, these results revealed a novel regulatory mechanism of SL biosynthesis and signaling in response to Pi starvation, providing genetic resources for improving plant architecture and nutrient-use efficiency in low-Pi environments.

A Crucial Role of GA-Regulated Flavonol Biosynthesis in Root Growth of Arabidopsis.

Flavonols have been demonstrated to play many important roles in plant growth, development, and communication with other organisms. Flavonol biosynthesis is spatiotemporally regulated by the subgroup 7 R2R3-MYB (SG7 MYB) transcription factors including MYB11/MYB12/MYB111. However, whether SG7-MYB activity is subject to post-translational regulation remains unclear. Here, we show that gibberellic acid (GA) inhibits flavonol biosynthesis via DELLA proteins in Arabidopsis. Protein-protein interaction analyses revealed that DELLAs (RGA and GAI) interacted with SG7 MYBs (MYB12 and MYB111) both in vitro and in vivo, leading to enhanced affinity of MYB binding to the promoter regions of key genes for flavonol biosynthesis and thus increasing their transcriptional levels. We observed that the level of auxin in the root tip was negatively correlated with root flavonol content. Furthermore, genetic assays showed that loss-of-function mutations in MYB12, which is predominantly expressed in roots, partially rescued the short-root phenotype of the GA-deficient mutant ga1-3 by increasing root meristem size and mature cell size. Consistent with these observations, exogenous application of the flavonol quercetin restored the root meristem size of myb12 ga1-3 to that of ga1-3. Taken together, our data elucidate a molecular mechanism by which GA promotes root growth by directly reducing flavonol biosynthesis.