Sucrose-activated TOR and phyA signaling alleviates shade-mediated inhibition of leaf development in Arabidopsis

To over-top the neighbors, shade-intolerant plants trigger shade avoidance responses with increasing the elongation of stem-like organs at the expense of leaf development. Shade-induced elongation growth has been extensively studied, while the regulatory mechanisms underlying shade-suppressed leaf development remain poorly understood. Here we find sucrose can rescue shade-inhibited leaf development, which requires the functions of phytochrome A (phyA) and TARGET OF RAPAMYCIN (TOR) complex. We confirm an interaction between PHYA and LETHAL WITH SEC THIRTEEN 8 (LST8), a component of TOR complex. Shade and phyA affect TOR activity, in turn TOR affects the protein stability of PHYA. The sucrose-TOR mediated accumulation of PHYA requires CONSTITUTIVELY PHOTOMORPHOGENIC1 (COP1)/SUPPRESSOR OF PHYA-105 (SPA) complex. Transcriptome analysis further reveals that sucrose and phyA regulate the expression of leaf development-related genes under shade, particularly those involved in cytokinin metabolism. Our work uncovers a critical role for TOR and PHYA in linking light signaling with cellular energy status, a mechanism that fine-tunes leaf development specifically under shade conditions.