A gas-and-brake mechanism of bHLH proteins modulates shade avoidance in Arabidopsis thaliana

Plants detect proximity of competitors through reduction in the ratio between red and far red light triggering the shade avoidance syndrome, which includes accelerated shoot elongation and early flowering. Shade avoidance is regulated through PHYTOCHROME INTERACTING FACTORs (PIFs), a group of bHLH transcription factors. Another (b)HLH protein, KIDARI (KDR), which is non-DNA-binding, was identified in de-etiolation studies and proposed to interact with LONG HYPOCOTYL IN FAR-RED 1 (HFR1), a (b)HLH protein that inhibits shade avoidance. Here we establish novel roles of KDR in regulating shade avoidance and investigate how KDR regulates the shade avoidance network. We show that KDR is a positive regulator of shade avoidance and interacts with several negative growth regulators. We identify novel interactors using a combination of yeast two-hybrid screening and dedicated confirmations with bimolecular fluorescence complementation. We demonstrate that KDR is translocated primarily to the nucleus when coexpressed with these newly discovered interactors. A genetic approach confirmed that several of these novel interactions are indeed functional to shade avoidance in , whereas we propose that KDR does not interact with HFR1 to regulate shade avoidance. Based on this, we propose that shade avoidance is regulated by a three-layered gas-and-brake mechanism of bHLH protein interactions, adding an additional layer of complexity to what was previously known.