Radial Growth in the Cycadales

The development of radial growth which leads to the pachycaulous form was investigated in eight of the 10 genera of the Cycadales; i.e., Ceratozamia, Cycas, Dioon, Encephalartos, Macrozamia, Microcycas, Stangeria, and Zamia. In all taxa, development of radial growth is essentially the same: a primary thickening meristem is differentiated in the stelar region of the cotyledonary node of the seedling at germination and produces derivatives mainly centrifugally. This primary thickening meristem (PTM) then differentiates acropetally and becomes continuous with the peripheral zone of the shoot apex. At first the PTM is a vertical cylinder, but as the seedling continues to grow into an adult plant, the PTM shows a more horizontal orientation (like an open umbrella) and produces the broad cortex. Secondary growth is by a vascular cambium which produces secondary xylem to the inside and secondary phloem to the outside. The broad pith originates from derivatives of the rib meristem of the massive shoot apex. The seedling and young plant is composed of a shortened shoot (i.e., no internodes) produced by the PTM and rib meristem, and a large fleshy primary root which results from a diffuse growth pattern. Individual cells in both the pith and cortex of the root divide. Their derivatives divide at right angles to the original division plane. Thus, quartets and even octets of cells are recognizable and can be traced to individual parent cells.