TRACHEID AND PIT ANATOMY VARY IN TANDEM IN A TALL SEQUOIADENDRON GIGANTEUM TREE

Across land plants there is a general pattern of xylem conduit diameters widening towards the stem base thus reducing the accumulation of hydraulic resistance as plants grow taller. In conifers, xylem conduits consist of cells with closed end-walls and water must flow through bordered pits imbedded in the side walls. As a consequence both cell size, which determines the numbers of walls that the conductive stream of water must cross, as well as the characteristics of the pits themselves, crucially affect total hydraulic resistance. Because both conduit size and pit features influence hydraulic resistance in tandem, we hypothesized that features of both should vary predictably with one another. To test this prediction we sampled a single tall (94.8 m) Sequoiadendron giganteum tree (giant sequoia), collecting wood samples from the most recent annual ring progressively downwards from the tree top to the base. We measured tracheid diameter and length, number of pits per tracheid, and the areas of pit apertures, tori, and margos. Tracheid diameter widened from treetop to base following a power law with an exponent (tracheid diameter-stem length slope) of approximately 0.20. A similar scaling exponent was found between tracheid length and distance from tree top. Additionally, pit aperture, torus, and margo areas all increased (again with a power of similar to 0.20) with distance from tree top, paralleling the observed variation in tracheid diameter and length. Pit density scaled isometrically with tracheid length. Within individual tracheids, total permeable area of pits, measured as the sum of the margo areas, scaled isometrically with lumen area. Given that pores of the margo membrane are believed to increase in parallel with membrane area, from a strictly anatomical perspective, our results support the interpretation that pit resistance remains a relatively constant fraction of total resistance along the hydraulic pathway.