Organization of the Vascular System in the Stems of Diplazium and Blechnum (Filicales)

Abstract

Detailed analysis of the three-dimensional vascular organization in species of Diplazium and Blechnum indicates that these ferns possess reticulate (dictyostelic) vascular systems that closely reflect the helical phyllotaxis of the shoot. In each species, the vascular pattern shows a specific relationship to the phyllotaxis, so that the phyllotactic fraction can be determined by examination of the number of cauline vascular bundles (meristeles) in cross section of the stem. The number of meristeles in a cross section equals the denominator of the phyllotactic fraction, i.e., the number of foliar orthostichies on the stem. The same numerical relationship also exists in the eusteles of seed plants between the number of axial (sympodial) stem bundles and the phyllotaxis. There is a further parallel between the three-dimensional reticulate pattern of fern dictyosteles and the reticulate patterns that characterize some herbaceous dicotyledons. However, the hypothesized separate origins of seed plant eusteles and fern dictyosteles from protostelic precursors preclude any direct homologies between these similar patterns. The parallel evolution of presumably more physiologically efficient reticulate systems in herbaceous seed plants and in ferns that have only a primary plant body is noteworthy. The similar relationships between the primary stem vascular patterns and phyllotaxy in both ferns and seed plants further emphasize the likely similarity of the morphogenetic events that occur at the shoot apex in these taxonomically disparate groups.