Gravitropism and gravimorphism during regeneration from protoplasts of the moss Ceratodon purpureus (Hedw.) Brid

Wild-type (WT) protonemata of the moss Ceratodon purpureus grow upwards in darkness (negative gravitropism), whereas protonemata of the mutant, wrong-way response (wwr-1) grow down. Since Ceratodon protoplasts regenerate to form new protonemata, we analyzed whether the direction of filament emergence was influenced by gravity (gravimorphism) and determined the cytological events that correlated with the onset of gravitropism in WT and wwr-1 filaments formed de novo. In the WT the direction of filament emergence appeared to be gravimorphic as more than 66% of the new filaments emerged above the horizontal. In contrast, the direction of filament emergence was random in wwr-1. Tip-growing cells of both genotypes became gravitropic within a total of one to two cell divisions. Gravitropic curvature in wwr-1 was opposite in direction to that of WT, and the timing of curvature was comparable, indicating that the wwr-1 mutation acts during the onset of gravitropic competence. In time-lapse studies of both genotypes, neither a plastid-free zone nor obvious and extensive plastid sedimentation characteristic of mature dark-grown protonemata was observed in the new filaments prior to gravitropic curvature. Thus, it appears that these latter two features are not required for gravitropism in new protonemal filaments from protoplasts.