Periodic root branching is influenced by light through an HY1-HY5-auxin pathway

The spacing of lateral roots (LRs) along the main root in plants is driven by an oscillatory signal, often referred to as the "root clock" that represents a pre-patterning mechanism that can be influenced by environmental signals. Light is an important environmental factor that has been previously reported to be capable of modulating the root clock, although the effect of light signaling on the LR pre-patterning has not yet been fully investigated. In this study, we reveal that light can activate the transcription of a photomorphogenic gene HY1 to maintain high frequency and amplitude of the oscillation signal, leading to the repetitive formation of pre-branch sites. By grafting and tissue-specific complementation experiments, we demonstrated that HY1 generated in the shoot or locally in xylem pole pericycle cells was sufficient to regulate LR branching. We further found that HY1 can induce the expression of HY5 and its homolog HYH, and act as a signalosome to modulate the intracellular localization and expression of auxin transporters, in turn promoting auxin accumulation in the oscillation zone to stimulate LR branching. These fundamental mechanistic insights improve our understanding of the molecular basis of light-controlled LR formation and provide a genetic interconnection between shoot- and root-derived signals in regulating periodic LR branching.