THE CONTROL OF THE ORIENTATION OF CELL DIVISIONS IN FERN GAMETOPHYTES

Time-lapse observations of filamentous fern gametophytes were used to evaluate whether the plane of cell division is referable to the plane of minimal surface area before and during the transition to two-dimensional growth. Cell dimensions of the apical cell were related to the length/width ratios associated with minimal area in the transverse plane vs. longitudinal plane, by modeling the apical cell as a hemisphere subtended by a cylinder. Our working hypothesis predicts that filamentous growth is perpetuated by an apical cell geometry that makes the transverse division plane the orientation of minimal surface area, whereas the transition to two-dimensional growth (longitudinal division of the apical cell) occurs once the longitudinal plane becomes the position of minimal surface area. The predictions of this hypothesis are fulfilled regardless of variations in light intensity and light quality, the presence of regulators of metabolism, or whether the experimental perturbation causes a corresponding selective inhibition of the transition to two-dimensional growth. Thus, the control of the plane of cell division in this system seems to depend on thermodynamic considerations of surface area. Furthermore, we favor the conclusion that the role of the genome in the transition to two-dimensional growth involves its influence on apical cell dimensions rather than the induction of specific genes for specific morphogenetic mRNAs.