ORIENTATION OF THE PLANE OF CELL DIVISION IN FERN GAMETOPHYTES: THE ROLES OF CELL SHAPE AND STRESS

Apical cells of Onoclea sensibilis L. protonemata were measured to determine areas of new walls which were formed during both transverse and longitudinal cell division. Actual wall areas were compared with calculated areas of hypothetical walls oriented in the opposite sense (i.e., an actual transverse wall compared with a hypothetical longitudinal wall, and the reverse). Among 87 out of 90 cells which were analyzed the actual walls had the least area. Thus, the minimal area hypothesis of cell partitioning accurately predicts wall orientation in this instance, although it appears, on other grounds, that the hypothesis does not furnish a plausible mechanism for wall orientation. The application of Lintilhac's concept of the orientation of cell walls in response to anisotropic stresses in the cell was explored. Photographs of apical cells during deplasmolysis indicated that unequal stresses might be generated in apical cells as a result of the osmotic distension of the elastic protoplast. It is concluded that the primary factor which determines the plane of cell division in the apical cell, and the transition from one- to two-dimensional growth, is the local pattern of stress which exists at the position of the nucleus at the time of onset of cell division and wall formation. Calculations of some geometrical properties of idealized model cells are interpreted to mean that the accuracy of the minimal area hypothesis results from a coincidence of its predictions with predictions of Lintilhac's hypothesis, and no causal significance is attributed to wall areas.