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Leaf-level anatomical variation is readily apparent within tall tree crowns, yet the relative importance of water and light availability in controlling this variation remains unclear. Sitka spruce (Picea sitchensis, (Bong.) Carr.) thrives in temperate rainforests of the Pacific Northwest, where it has historically reached heights >100m, despite rarely living more than 400 years alongside redwoods that are five times older. We examined leaves of trees up to 97m tall using a combination of transverse sections, longitudinal sections, epidermal imprints and whole-leaf measurements to explore the combined effects of water stress and light availability on leaf development in P. sitchensis. In contrast to the situation in tall Cupressaceae, light availability--not hydraulic limitation--is the primary ecological driver of leaf-level anatomical variation in P. sitchensis. While height-associated decreases in leaf length and mesoporosity are best explained by hydrostatic constraints on leaf elongation, the majority of anatomical traits we measured reflect acclimation to light availability, including increases in leaf width and vascular tissue areas in the brightest parts of the crown. Along with these changes, the appearance of abaxial stomata in the bright upper crown, and the arrangement of mesophyll in uniseriate, transverse plates--with radially arranged apoplastic pathways leading directly to stomata before bridging them with a V-shaped cell--may enhance gas exchange and hydraulic conductivity. This suite of leaf traits suggests an adaptive strategy that maximizes photosynthesis at the expense of water-stress tolerance. Anatomical investigations spanning the height gradient in tall tree crowns build our understanding of mechanisms underlying among-species variation in growth rates, life spans, and potential responses to climate change. [ABSTRACT FROM AUTHOR]

We examined changes in branch hydraulic, leaf structure and gas exchange properties in coast redwood ( Sequoia sempervirens) and giant sequoia ( Sequoiadendron giganteum) trees of different sizes. Leaf-specific hydraulic conductivity ( kL) increased with height in S. sempervirens but not in S. giganteum, while xylem cavitation resistance increased with height in both species. Despite hydraulic adjustments, leaf mass per unit area (LMA) and leaf carbon isotope ratios ( δ13C) increased, and maximum mass-based stomatal conductance ( gmass) and photosynthesis ( Amass) decreased with height in both species. As a result, both Amass and gmass were negatively correlated with branch hydraulic properties in S. sempervirens and uncorrelated in S. giganteum. In addition, Amass and gmass were negatively correlated with LMA in both species, which we attributed to the effects of decreasing leaf internal CO2 conductance ( gi). Species-level differences in wood density, LMA and area-based gas exchange capacity constrained other structural and physiological properties, with S. sempervirens exhibiting increased branch water transport efficiency and S. giganteum exhibiting increased leaf-level water-use efficiency with increasing height. Our results reveal different adaptive strategies for the two redwoods that help them compensate for constraints associated with growing taller, and reflect contrasting environmental conditions each species faces in its native habitat. [ABSTRACT FROM AUTHOR]

Trees grow tall where resources are abundant, stresses are minor, and competition for light places a premium on height growth. The height to which trees can grow and the biophysical determinants of maximum height are poorly understood. Some models predict heights of up to 120?m in the absence of mechanical damage, but there are historical accounts of taller trees. Current hypotheses of height limitation focus on increasing water transport constraints in taller trees and the resulting reductions in leaf photosynthesis. We studied redwoods (Sequoia sempervirens), including the tallest known tree on Earth (112.7?m), in wet temperate forests of northern California. Our regression analyses of height gradients in leaf functional characteristics estimate a maximum tree height of 122-130?m barring mechanical damage, similar to the tallest recorded trees of the past. As trees grow taller, increasing leaf water stress due to gravity and path length resistance may ultimately limit leaf expansion and photosynthesis for further height growth, even with ample soil moisture. [ABSTRACT FROM AUTHOR]

Here we respond to the communication in American Journal of Botany (96: 542-544 in this issue) by Neting, who proposes several ways in which our paper "The Limits to Tree Height" (Nature 428: 851-854) may have erred in estimating the biophysical limits to height growth in Sequoia sempervirens. We first explain that because embolism repair requires long time periods and is generally incomplete, xylem vulnerability characteristics offer a sound basis for estimating performance limits in woody plants. We reaffirm our earlier use of vertical gradients of foliar carbon isotope composition with new data for S. sempervirens. We support these arguments with reference to studies in other tree species. We take exception with Netting's view that the turgor pressure-cell expansion relationship for Zea mays is applicable to S. sempervirens. Finally, we second Netting's call for more work on carbon allocation vis a vis height growth limits. [ABSTRACT FROM AUTHOR]