The localization of phototropin to the plasma membrane defines a cold-sensing compartment in Marchantia polymorpha

Plant cells perceive cold temperatures and initiate cellular responses to protect themselves against cold stress, but which cellular compartment mediates cold sensing has been unknown. Chloroplasts change their position in response to cold to optimize photosynthesis in plants in a process triggered by the blue-light photoreceptor phototropin (phot), which thus acts as a cold-sensing molecule. However, phot in plant cells is present in multiple cellular compartments, including the plasma membrane (PM), cytosol, Golgi apparatus, and chloroplast periphery, making it unclear where phot perceives cold and activates this cold-avoidance response. Here, we produced genetically encoded and modified variants of phot that localize only to the cytosol or the PM and determined that only PM-associated phot induced cold avoidance in the liverwort Marchantia polymorpha. These results indicate that the phot localized to the PM constitutes a cellular compartment for cold sensing in plants.