title: The hydrostatic gradient, not light availability, drives height-related variation in Sequoia sempervirens (Cupressaceae) leaf anatomy.
abstract: OBJECTIVE
  Leaves at the tops of most trees are smaller, thicker, and in many other ways different from leaves on the lowermost branches. This height-related variation in leaf structure has been explained as acclimation to differing light environments and, alternatively, as a consequence of hydrostatic, gravitational constraints on turgor pressure that reduce leaf expansion. •
  
  
  METHODS
  To separate hydrostatic effects from those of light availability, we used anatomical analysis of height-paired samples from the inner and outer tree crowns of tall redwoods (Sequoia sempervirens). •
  
  
  RESULTS
  Height above the ground correlates much more strongly with leaf anatomy than does light availability. Leaf length, width, and mesophyll porosity all decrease linearly with height and help explain increases in leaf-mass-to-area ratio and decreases in both photosynthetic capacity and internal gas-phase conductance with increasing height. Two functional traits-leaf thickness and transfusion tissue-also increase with height and may improve water-stress tolerance. Transfusion tissue area increases enough that whole-leaf vascular volume does not change significantly with height in most trees. Transfusion tracheids become deformed with height, suggesting they may collapse under water stress and act as a hydraulic buffer that improves leaf water status and reduces the likelihood of xylem dysfunction. •
  
  
  CONCLUSIONS
  That such variation in leaf structure may be caused more by gravity than by light calls into question use of the terms "sun" and "shade" to describe leaves at the tops and bottoms of tall tree crowns.
authors: Department of Biological Sciences, Humboldt State University, Arcata, California 95521 USA.
authors: Alana R
authors: AR
authors: Oldham
authors: Stephen C
authors: SC
authors: Sillett
authors: Alexandru M F
authors: AM
authors: Tomescu
authors: George W
authors: GW
authors: Koch