The leaf area to sapwood area ratio (A :A) of trees has been hypothesized to decrease as trees become older and taller. Theory suggests that A :A must decrease to maintain leaf-specific hydraulic sufficiency as path length, gravity, and tortuosity constrain whole-plant hydraulic conductance. We tested the hypothesis that A :A declines with tree height. Whole-tree A :A was measured on 15 individuals of Douglas-fir (Pseudotsuga menziesii var. menziesii) ranging in height from 13 to 62 m (aged 20-450 years). A :A declined substantially as height increased (P=0.02). Our test of the hypothesis that A :A declines with tree height was extended using a combination of original and published data on nine species across a range of maximum heights and climates. Meta-analysis of 13 whole-tree studies revealed a consistent and significant reduction in A :A with increasing height (P<0.05). However, two species (Picea abies and Abies balsamea) exhibited an increase in A :A with height, although the reason for this is not clear. The slope of the relationship between A :A and tree height (ΔA :A/Δh) was unrelated to mean annual precipitation. Maximum potential height was positively correlated with ΔA :A/Δh. The decrease in A :A with increasing tree size that we observed in the majority of species may be a homeostatic mechanism that partially compensates for decreased hydraulic conductance as trees grow in height.
The hydraulic limitation hypothesis proposes that (1) reduced growth in taller trees is caused by decreased photosynthesis resulting from a decrease in hydraulic conductance promoted by a longer root-to-leaf flow path, and (2) this mechanism reduces stand productivity after canopy closure. This hypothesis was tested by comparing the phys-
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