Phytochrome B regulates cortical microtubule arrangement to control cotyledon polar expansion by repressing <i>LONGIFOLIA</i>s

Light promotes the expansion and controls the directionality of expansion in cotyledons, transforming small oval cotyledons into larger orbicular shapes. However, the cellular basis underlying this polar expansion remains unclear. We report that cotyledon polar expansion in Arabidopsis (Arabidopsis thaliana) is primarily associated with the polar expansion of pavement cells, rather than with polar cell proliferation. Phytochrome B (phyB) promotes this polar expansion by inhibiting PHYTOCHROME INTERACTING FACTORs (PIFs), which normally suppress expansion and inversely regulate its directionality. PIFs exert their control over directionality partly through the activation of their target genes, LONGIFOLIAs (LNGs). At the cellular level, phyB decreases the number of transversely arranged cortical microtubules, while increasing the number of longitudinally arranged microtubules. This phyB-induced change in microtubule arrangement would strengthen transverse expansion while weakening longitudinal expansion. In contrast, PIFs regulate microtubule arrangements in the opposite manner. Downstream of the phyB-PIF pathway, LNGs preferentially increase transversely arranged cortical microtubules. Overall, our data support that the regulation of cortical microtubule orientation by the phyB-PIF-LNG pathway underlies how phyB weakens longitudinal expansion relative to transverse expansion while promoting pavement cell expansion to make orbicular cotyledons in the light.