SENSITIVE TO FREEZING2 is crucial for growth of Marchantia polymorpha under acidic conditions.

Land plants have evolved many systems to adapt to a wide range of environmental stresses. In seed plants, oligogalactolipid synthesis is involved in tolerance to freezing and dehydration, but it has not been analyzed in non-vascular plants. Here we analyzed trigalactosyldiacylglycerol (TGDG) synthesis in Marchantia polymorpha. TGDG is synthesized by galactolipid: galactolipid galactosyltransferase [GGGT; SENSITIVE TO FREEZING2 (SFR2) in Arabidopsis]. We analyzed the subcellular localization and GGGT activity of two M. polymorpha SFR2 homologs (MpGGGT1 and MpGGGT2, each as a GFP-fusion protein) using a transient expression system in Nicotiana benthamiana leaves and found that MpGGGT1-GFP localized in the chloroplast envelope membrane. We produced mutants Mpgggt1 and Mpgggt2 and found that TGDG did not accumulate in Mpgggt1 upon treatment of the thallus with acetic acid. Moreover, growth of Mpgggt1 mutants was impaired by acetic acid treatment. Microscopy revealed that the acetic acid treatment of M. polymorpha plants damaged intracellular membranes. The fact that the effect was similar for wild-type and Mpgggt1 plants suggested that MpGGGT has a role in recovery from damage. These results indicate that MpGGGT plays a crucial role in M. polymorpha growth under conditions of acid stress, which may have been encountered during the ancient terrestrial colonization of plants.