Brassinosteroids Influence Arabidopsis Hypocotyl Graviresponses Through Changes In Mannans And Cellulose

The force of gravity is a constant environmental factor. Plant shoots respond to gravity through negative gravitropism and gravity resistance. These responses are essential for plants to direct the growth of aerial organs away from the soil surface after germination and to keep an upright posture above ground. We took advantage of the effect of brassinosteroids on the two types of graviresponses in Arabidopsis thaliana hypocotyls to disentangle functions of cell wall polymers during etiolated shoot growth. The ability of etiolated Arabidopsis seedlings to grow upwards was suppressed in the presence of 24-epibrassinolide (EBL) but enhanced in the presence of brassinazole (BRZ), an inhibitor of brassinosteroid biosynthesis. These effects were accompanied by changes in cell wall mechanics and composition. Cell wall biochemical analyses and confocal microscopy of the cellulose-specific pontamine S4B dye revealed that the EBL and BRZ treatments correlated with changes in cellulose fibre organization and mannan content. Indeed, a longitudinal re-orientation of cellulose fibres supported upright growth whereas the presence of mannans reduced gravitropic bending. The negative effect of mannans on gravitropism is a new function for this class of hemicelluloses, highlighting evolutionary adaptations by which aquatic ancestors of terrestrial plants colonized land.