The twisted dwarf1 (twd1) mutant displays a pleiotropic developmental phenotype resembling atpgp1-1/atpgp19-1 (atmdr1-1) double mutants. (A) Phenotype of soil-grown wild-type (left) and twd1-1 (right) plants at maturity. Bars, 5 cm. (B) Siliques of wild-type, atpgp1-1, atpgp19-1, atpgp1-1/atpgp19-1, and twd1-1 (from left to right) plants showing disoriented growth behavior. Bars, 1 cm. (C) Light grown seedlings 5 d after germination. From left to right: wild-type, atpgp1-1, atmdr1-1 (atpgp19-1), atmdr1-1/atpgp1-1, and twd1-1. Bar, 1 cm. (D) Rosette leaves of wild-type, atpgp1-1, atpgp19-1, atpgp1-1/atpgp19-1, and twd1-1 (from left to right) plants. Double mutant and twd1-1 show strongly reduced leaf expansion and strong epinastic growth behavior. Bar, 1 cm. (E) Dark-grown seedlings of atpgp1-1/atpgp19-1 (atmdr1-1), and twd1-1 plants have longer roots. Seedlings were grown on plate in darkness and root lengths. Root lengths were measured with a ruler (>10 seedlings) after 8 d and are presented as means plus SDs. Plant growth being statistically different (Mann-Whitney U test, p > 0.05) compared with wild-type control plants is indicated by an asterisks. (F) Seedlings of atpgp1-1/atpgp19-1 (atmdr1-1) and twd1-1 plants have longer hypocotyls. Seedlings were grown on plate in darkness or continuous white light. Hypocotyl lengths were measured with a ruler (> 10 seedlings) after 8 d and are presented as means plus SDs. Plant growth being statistically different (Mann-Whitney U test, p > 0.05) compared with wild-type control plants is indicated by an asterisks. (G) Phenotype of soil-grown plants after 40 d of culture. Top panel from left to right: wild-type, atpgp1-1, atpgp19-1 (atmdr1-1), bottom panel: atpgp1-1/atpgp19-1 (atmdr1-1) double mutant, twd1-1. Bar, 5 cm. (H) Reduced apical dominance in atpgp1-1/atpgp19-1 (atmdr1-1) and twd1-1 plants. Plants (left panel from top to bottom: wild-type, atpgp1-1, and atpgp19-1 (atmdr1-1); right panel from left to right: atpgp1-1/atpgp19-1 (atmdr1-1) and twd1-1) were cultured on soil for 70 d. Note the size bars differ between left and right panels.