Developmental anatomy of the three-dimensional leaf ofBotrychium ternatum (Thunb.) Sw.

The ophioglossaceous leaf exhibits a unique morphology, three-dimensional constitution. This examination clarified the developmental manner of the leaf ofBotrychium ternatum (Thunb.) Sw. The leaf primordium develops as an ordinary appendicular leaf which shows a typical hyponastic growth curvature, though it soon takes a conical shape with a tetrahedral apical cell. The leaf primordium forms a sporophyll primordium first, then forms vegetative primary pinnae acting as the trophophyll primordium. The sporophyll primordium also forms sporogenous primary pinnae. The sporophyll initiation begins with establishment of a new apical cell (sporophyll apical cell) near the original leaf apical cell. Through activity of the sporophyll apical cell most of the sporophyll primordium is formed later. In contrast, the vegetative or sporogenous primary pinna begins to develop as a low mound of surface cells on the trophophyll or sporophyll primordium respectively. The apical cell is later established on the summit of each pinna primordium. In the developmental point of view, the sporophyll primordium is not equivalent to the primary pinnae on the trophophyll primordium. The sporophyll may not represent two fused basal pinnae of a leaf, but represents an organ independent of and equivalent to the trophophyll.