Proline Affects Flowering Time in Arabidopsis by Modulating FLC Expression: A Clue of Epigenetic Regulation?

The recent finding that proline-induced root elongation is mediated by reactive oxygen species (ROS) prompted us to re-evaluate other developmental processes modulated by proline, such as flowering time. By controlling the cellular redox status and the ROS distribution, proline could potentially affect the expression of transcriptional factors subjected to epigenetic regulation, such as FLOWERING LOCUS C (FLC). Accordingly, we investigated the effect of proline on flowering time in more detail by analyzing the relative expression of the main flowering time genes in p5cs1 p5cs2/P5CS2 proline-deficient mutants and found a significant upregulation of FLC expression. Moreover, proline-deficient mutants exhibited an adult vegetative phase shorter than wild-type samples, with a trichome distribution reminiscent of plants with high FLC expression. In addition, the vernalization-induced downregulation of FLC abolished the flowering delay of p5cs1 p5cs2/P5CS2, and mutants homozygous for p5cs1 and flc-7 and heterozygous for P5CS2 flowered as early as the flc-7 parental mutant, indicating that FLC acts downstream of P5CS1/P5CS2 and is necessary for proline-modulated flowering. The overall data indicate that the effects of proline on flowering time are mediated by FLC.