Tracheid buckling buys time, foliar water uptake pays it back: Coordination of leaf structure and function in tall redwood trees.

Tracheid buckling may protect leaves in the dynamic environments of forest canopies, where rapid intensifications of evaporative demand, such as those brought on by changes in light availability, can result in sudden increases in transpiration rate. While treetop leaves function in reliably direct light, leaves below the upper crown must tolerate rapid, thermally driven increases in evaporative demand. Using synchrotron-based X-ray microtomography, we visualized impacts of experimentally induced water stress and subsequent fogging on living cells in redwood leaves, adding ecological and functional context through crown-wide explorations of variation in leaf physiology and microclimate.

Shoot dimorphism enables Sequoia sempervirens to separate requirements for foliar water uptake and photosynthesis.

Premise: Trees in wet forests often have features that prevent water films from covering stomata and inhibiting gas exchange, while many trees in drier environments use foliar water uptake to reduce water stress. In forests with both wet and dry seasons, evergreen trees would benefit from producing leaves capable of balancing rainy-season photosynthesis with summertime water absorption.

Methods: Using samples collected from across the vertical gradient in tall redwood (Sequoia sempervirens) crowns, we estimated tree-level foliar water uptake and employed physics-based causative modeling to identify key functional traits that determine uptake potential by setting hydraulic resistance.

Within-crown plasticity in leaf traits among the tallest conifers.

Premise Of The Study: Leaves are the sites of greatest water stress in trees and a key means of acclimation to the environment. We considered phenotypic plasticity of Pseudotsuga menziesii leaves in their ecological context, exploring responsiveness to natural gradients in water stress (indicated by sample height) and light availability (measured from hemispherical photos) to understand how leaf structure is controlled by abiotic factors in tall tree crowns.

Methods: After measuring anatomy, morphology, and carbon isotope composition (δ C) of leaves throughout crowns of P.

Leaf acclimation to light availability supports rapid growth in tall Picea sitchensis trees.

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.

Phenotypic plasticity of leaves enhances water-stress tolerance and promotes hydraulic conductivity in a tall conifer.

Premise Of The Study: Leaves respond to environmental signals and acclimate to local conditions until their ecological limits are reached. Understanding the relationships between anatomical variation in leaves and the availability of water and light improves our ability to predict ecosystem-level impacts of foliar response to climate change, as it expands our knowledge of tree physiology.

Methods: We examined foliar anatomy and morphology of the largest plant species, Sequoiadendron giganteum, from leafy shoot samples collected throughout crowns of trees up to 95 m tall and assessed the functionality of within-crown variation with a novel drought/recovery experiment.

Growth maximization trumps maintenance of leaf conductance in the tallest angiosperm.

Structural and physiological changes that occur as trees grow taller are associated with increased hydraulic constraints on leaf gas exchange, yet it is unclear if leaf-level constraints influence whole-tree growth as trees approach their maximum size. We examined variation in leaf physiology, leaf area to sapwood area ratio (L/S), and annual aboveground growth across a range of tree heights in Eucalyptus regnans. Leaf photosynthetic capacity did not differ among upper crown leaves of individuals 61.

Millennium-scale crossdating and inter-annual climate sensitivities of standing California redwoods.

Extremely decay-resistant wood and fire-resistant bark allow California's redwoods to accumulate millennia of annual growth rings that can be useful in biological research. Whereas tree rings of Sequoiadendron giganteum (SEGI) helped formalize the study of dendrochronology and the principle of crossdating, those of Sequoia sempervirens (SESE) have proven much more difficult to decipher, greatly limiting dendroclimatic and other investigations of this species. We overcame these problems by climbing standing trees and coring trunks at multiple heights in 14 old-growth forest locations across California.

Structural development of redwood branches and its effects on wood growth.

Redwood branches provide all the carbohydrates for the most carbon-heavy forests on Earth, and recent whole-tree measurements have quantified trunk growth rates associated with complete branch inventories. Providing all of a tree's photosynthetic capacity, branches represent an increasing proportion of total aboveground wood production as trees enlarge. To examine branch development and its effects on wood volume growth, we dissected 31 branches from eight Sequoia sempervirens (D.

The hydrostatic gradient, not light availability, drives height-related variation in Sequoia sempervirens (Cupressaceae) leaf anatomy.

Premise Of The Study: 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).

Effects of height on treetop transpiration and stomatal conductance in coast redwood (Sequoia sempervirens).

Treetops become increasingly constrained by gravity-induced water stress as they approach maximum height. Here we examine the effects of height on seasonal and diurnal sap flow dynamics at the tops of 12 unsuppressed Sequoia sempervirens (D. Don) Endl.

Physiological consequences of height-related morphological variation in Sequoia sempervirens foliage.

This study examined relationships between foliar morphology and gas exchange characteristics as they vary with height within and among crowns of Sequoia sempervirens D. Don trees ranging from 29 to 113 m in height. Shoot mass:area (SMA) ratio increased with height and was less responsive to changes in light availability as height increased, suggesting a transition from light to water relations as the primary determinant of morphology with increasing height.

A response to: Limitations within "The Limits to Tree Height".

Here we respond to the communication in American Journal of Botany (96: 542-544 in this issue) by Netting, 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.

Hydrostatic constraints on morphological exploitation of light in tall Sequoia sempervirens trees.

We studied changes in morphological and physiological characteristics of leaves and shoots along a height gradient in Sequoia sempervirens, the tallest tree species on Earth, to investigate whether morphological and physiological acclimation to the vertical light gradient was constrained by hydrostatic limitation in the upper crown. Bulk leaf water potential (Psi) decreased linearly and light availability increased exponentially with increasing height in the crown. During the wet season, Psi was lower in the outer than inner crown.

The limits to tree height.

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.

Effects of tree crown structure on biomass of the epiphytic fern Polypodium scouleri (Polypodiaceae) in redwood forests.

Redwood forests contain some of the largest and most structurally complex trees on Earth. The most abundant vascular epiphyte in these forests is the fern Polypodium scouleri (Polypodiaceae). We measured dimensions of all 765 P.