Cell biology of bordered-pit formation in balsam-fir trees

A mature bordered pit in secondary xylem of Pinaceae comprises a circular border of secondary-wall material that protrudes into the tracheid lumen and is punctuated by a centralized aperture through which sap flows. The overarching border encloses a pit chamber within which is a membrane, or diaphragm, consisting of a central torus and margo strands. Bordered-pit pairs are abundantly present in all woods, and their membranes serve as swinging-diaphragm check valves regulating sap flow between adjoining tracheary elements, simultaneously trapping emboli and particulates in water as it moves from roots to leaves. The cell biology of bordered-pit formation in cambial derivatives of Abies balsamea (L.) Mill. was investigated by light and scanning electron microscopy during early stages of cellular differentiation of cambial derivatives into secondary xylem tracheids. A bordered-pit template (BPT), a bordered-pit organelle (BPO), a bordered-pit basal disk, and additional novel structures were found to be associated with bordered-pit formation. Evidence was found that the membrane does not comprise residual compound middle lamella; rather, the membrane forms de novo as BPO remnants. A cell-biology model and new terminology are introduced to explain how BPTs, BPOs, and basal disks contribute to successive stages in formation of bordered-pit pairs.