XYLEM ANATOMY,WATER FLOW,AND HYDRAULIC CONDUCTANCE IN THE FERN CYRTOMIUM FALCATUM

Xylem anatomy and water relations were studied in holly fern (Cyrtomium falcatum, Aspidiaceae) to determine the details of the pathway for water flow through an entire plant and the influence of tracheid number and lumen diameter on water flow. Each leaf has two adaxial traces and an abaxial trace, which are supplied by diarch adventitious roots attached to the dictyostele of the rhizome near the leaf base. Anatomical observations and dye experiments showed that each adaxial bundle vascularizes the approximately seven pinnae on its side of a leaf. An abaxial bundle is intermittently connected to an adaxial bundle as well as other abaxial bundles, forming a minor vascular pathway between the bundles of the leaf axis. Changes in both number and diameter of tracheids result in an acropetal decrease in hydraulic conductance per unit length along the rachis, although tracheid number locally increases when the trace for a pinna is produced in an adaxial bundle. Water flow was determined from the transpiration distal to the point in question or by forcing a solution through an axis with applied pressure. The water potential gradient along the plant axis was quite constant, indicating that hydraulic conductance per unit length varied with leaf area to be supplied. About 40% of the overall water potential drop occurred from the rachis into the pinnae, which reflected factors controlling water potential gradients in the lamina and not a very low conductance in the petiolule xylem. Hydraulic conductances calculated using the Hagen-Poiseuille equation and tracheid diameters were generally double those of measured conductances. Since the values tended to vary by a constant factor, tracheid number and diameter may largely control water flow in the xylem.