8 USA 9 10 keywords: mechanosensitive channel, Arabidopsis, mitochondria, chloroplast, reactive oxygen 11 species 12 13 2 ABSTRACT 14 15 Plant development requires communication on many levels, including between cells and between 16 organelles within a cell. For example, mitochondria and plastids have been proposed to be sensors 17 of environmental stress and to coordinate their responses. Here we present evidence for 18 communication between mitochondria and chloroplasts during leaf and root development, based 19 on genetic and physical interactions between three Mechanosensitive channels of Small 20 conductance-Like (MSL) proteins from Arabidopsis thaliana. MSL proteins are Arabidopsis 21 homologs of the bacterial Mechanosensitive channel of Small conductance (MscS), which relieves 22 cellular osmotic pressure to protect against lysis during hypoosmotic shock. MSL1 localizes to the 23 inner mitochondrial membrane, while MSL2 and MSL3 localize to the inner plastid membrane 24and are required to maintain plastid osmotic homeostasis during normal growth and development.
25In this study, we characterized the phenotypic effect of a genetic lesion in MSL1, both in wild type 26 and in msl2 msl3 mutant backgrounds. msl1 single mutants appear wild type for all phenotypes 27 examined. The characteristic leaf rumpling in msl2 msl3 double mutants was exacerbated in the 28 msl1 msl2 msl3 triple mutant. However, the introduction of the msl1 lesion into the msl2 msl3 29 mutant background suppressed other msl2 msl3 mutant phenotypes, including ectopic callus 30 formation, accumulation of superoxide and hydrogen peroxide in the shoot apical meristem, 31 decreased root length, and reduced number of lateral roots. All these phenotypes could be 32 recovered by molecular complementation with a transgene containing a wild type version of MSL1.
33In yeast-based interaction studies, MSL1 interacted with itself, but not with MSL2 or MSL3. These 34 results establish that the abnormalities observed in msl2 msl3 double mutants is partially dependent 35 on the presence of functional MSL1 and suggest a possible role for communication between plastid 36 and mitochondria in seedling development.37 38 39 40 Plastids and mitochondria are found in almost every plant cell and are involved in all aspects of 41 plant biology. In plants, as in animals, mitochondria are involved in multiple cellular processes, 42 including cellular respiration and co-enzyme synthesis (Schertl and Braun, 2014; Rébeillé et al., 43 2007). Plastids are responsible for photosynthesis and a range of other biosynthetic reactions-44 including the production of starch, some amino acids, fatty acids and lipids, pigments, hormones 45 and volatiles (Neuhaus:2000eh; Rolland et al., 2018). Some plastids play a unique role in plant 46 biology: amyloplasts in the root tip and the shoot endodermis are essential for gravity response 47 (Toyota:2013fe; Su et al., 2017). A recent report argues that plastids of the leaf epidermis can serve 48 as stress sensors (Beltrán et al., 2018). Whi...