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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 (δ13C) of leaves throughout crowns of P. menziesii >90 m tall, we compared structural plasticity of leaves among the three tallest conifer species using equivalent data from past work on Sequoia sempervirens and Picea sitchensis. Key results: Leaf mass per projected area (LMA) and δ13C increased and mesoporosity (airspace/area) decreased along the water‐stress gradient, while light did not play a detectable role in leaf development. Overall, leaves of P. menziesii were far less phenotypically responsive to within‐crown abiotic gradients than either P. sitchensis, whose leaves responded strongly to light availability, or S. sempervirens, whose leaves responded equally strongly to water stress. Conclusions: P. menziesii maintain remarkably consistent leaf structure despite pronounced vertical gradients in abiotic factors. Contrasting patterns of leaf structural plasticity underlie divergent ecological strategies of the three tallest conifer species, which coexist in Californian rainforests. [ABSTRACT FROM AUTHOR]

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 ab ility 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, Sequoiadendror giganteum, from leafy shoot samples collected th roughout crowns of trees up to 95 m tall and assessed the functionality of with in-crown variation with a novel drought/recovery experiment. KEY RESULTS: We found phenotypic variation in response to water availability in 13 anatomical traits of Sequoiadendron leaves. Shoot expansion was constrained by the hydrostatic gradient of maximum water potential, while functional traits supporting succulence and toughness were associated with sites of peak hydraulic limitation. Water-stress tolerance in experimental shoots increased dramatically with height. CONCLUSION: We propose a heat-sink function for transfusion tissue and uncover a suite of traits suggesting rapid hydraulic throughput and flexibility in water-stress tolerance investments as strategies that help this montane species reach such enormous size. Responses to water stress alter the amount of carbon stored in foliage and the rate of the eventual release of carbon. [ABSTRACT FROM AUTHOR]

The constraint on vertical water transport is considered an important factor limiting height growth and maximum attainable height of trees. Here, we show evidence of foliar water storage as a mechanism that could partially compensate for this constraint in Sequoia sempervirens, the tallest species., We measured hydraulic and morpho-anatomical characteristics of foliated shoots of tall S. sempervirens trees near the wet, northern and dry, southern limits of its geographic distribution in California, USA., The ability to store water (hydraulic capacitance) and saturated water content (leaf succulence) of foliage both increased with height and light availability, maintaining tolerance of leaves to water stress (bulk-leaf water potential at turgor loss) constant relative to height., Transverse-sectional area of water-storing, transfusion tissue in leaves increased with height, while the area of xylem tissue decreased, indicating increasing allocation to water storage and decreasing reliance on water transport from roots., Treetop leaves of S. sempervirens absorb moisture via leaf surfaces and have potential to store more than five times the daily transpirational demand. Thus, foliar water storage may be an important adaptation that helps maintain physiological function of treetop leaves and hydraulic status of the crown, allowing this species to partially compensate for hydraulic constraints and sustain turgor for both photosynthesis and height growth. [ABSTRACT FROM AUTHOR]