Homoplastic occurrence of perforated pit membranes and torus-bearing pit membranes in ancestral angiosperms as observed by field-emission scanning electron microscopy

Recent studies demonstrated that perforated pit membranes (i.e., pit membranes with a large opening in their central portion) are commonly present between wood fibers in core eudicots. It is unclear whether this type of pit membranes might also occur in ancestral angiosperms. Therefore, structure of interfiber pit membranes was examined by field-emission scanning electron microscopy in nine species representing seven families that are located at more ancestral position than core eudicots. We found perforated pit membranes in three of the nine species. Our observations indicate that perforated pit membranes are relatively common even in ancestral groups of angiosperms. In the non-perforated pit membranes of the other six species, we found a range of structural variations. Thin-walled pit membranes without apparent intercellular layers were always found in three of the six species and the porosity of sheet-like pit membranes differed among the three species. Unlike the thin-walled pit membranes, interfiber pit membranes of Buxus microphylla var. japonica were thick-walled with obvious intercellular layers, and in Schisandra chinensis, we often observed torus-bearing pit membranes. Such variations in layered structure of pit membranes and homoplastic occurrence of torus-bearing pit membranes have not yet been reported for ancestral angiosperms. Our observations indicate that the structure of interfiber pit membranes might be more complicated than previous studies might suggest.