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  1. Abstract

    Within vascular plants, the partitioning of hydraulic resistance along the soil‐to‐leaf continuum affects transpiration and its response to environmental conditions. In trees, the fractional contribution of leaf hydraulic resistance (Rleaf) to total soil‐to‐leaf hydraulic resistance (Rtotal), or fRleaf(=Rleaf/Rtotal), is thought to be large, but this has not been tested comprehensively. We compiled a multibiome data set of fRleafusing new and previously published measurements of pressure differences within trees in situ. Across 80 samples, fRleafaveraged 0.51 (95% confidence interval [CI] = 0.46−0.57) and it declined with tree height. We also used the allometric relationship between field‐based measurements of soil‐to‐leaf hydraulic conductance and laboratory‐based measurements of leaf hydraulic conductance to compute the average fRleaffor 19 tree samples, which was 0.40 (95% CI = 0.29−0.56). The in situ technique produces a more accurate descriptor of fRleafbecause it accounts for dynamic leaf hydraulic conductance. Both approaches demonstrate the outsized role of leaves in controlling tree hydrodynamics. A larger fRleafmay help stems from loss of hydraulic conductance. Thus, the decline in fRleafwith tree height would contribute to greater drought vulnerability in taller trees and potentially to their observed disproportionate drought mortality.

     
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  3. Summary

    Understanding the pronounced seasonal and spatial variation in leaf carboxylation capacity (Vc,max) is critical for determining terrestrial carbon cycling in tropical forests. However, an efficient and scalable approach for predictingVc,maxis still lacking.

    Here the ability of leaf spectroscopy for rapid estimation ofVc,maxwas tested.Vc,maxwas estimated using traditional gas exchange methods, and measured reflectance spectra and leaf age in leaves sampled from tropical forests in Panama and Brazil. These data were used to build a model to predictVc,maxfrom leaf spectra.

    The results demonstrated that leaf spectroscopy accurately predictsVc,maxof mature leaves in Panamanian tropical forests (R2 = 0.90). However, this single‐age model required recalibration when applied to broader leaf demographic classes (i.e. immature leaves). Combined use of spectroscopy models forVc,maxand leaf age enabled construction of theVc,max–age relationship solely from leaf spectra, which agreed with field observations. This suggests that the spectroscopy technique can capture the seasonal variability inVc,max, assuming sufficient sampling across diverse species, leaf ages and canopy environments.

    This finding will aid development of remote sensing approaches that can be used to characterizeVc,maxin moist tropical forests and enable an efficient means to parameterize and evaluate terrestrial biosphere models.

     
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  4. Abstract

    Forest leaf area has enormous leverage on the carbon cycle because it mediates both forest productivity and resilience to climate extremes. Despite widespread evidence that trees are capable of adjusting to changes in environment across both space and time through modifying carbon allocation to leaves, many vegetation models use fixed carbon allocation schemes independent of environment, which introduces large uncertainties into predictions of future forest responses to atmospheric CO2fertilization and anthropogenic climate change. Here, we develop an optimization‐based model, whereby tree carbon allocation to leaves is an emergent property of environment and plant hydraulic traits. Using a combination of meta‐analysis, observational datasets, and model predictions, we find strong evidence that optimal hydraulic–carbon coupling explains observed patterns in leaf allocation across large environmental and CO2concentration gradients. Furthermore, testing the sensitivity of leaf allocation strategy to a diversity in hydraulic and economic spectrum physiological traits, we show that plant hydraulic traits in particular have an enormous impact on the global change response of forest leaf area. Our results provide a rigorous theoretical underpinning for improving carbon cycle predictions through advancing model predictions of leaf area, and underscore that tree‐level carbon allocation to leaves should be derived from first principles using mechanistic plant hydraulic processes in the next generation of vegetation models.

     
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  5. Summary

    Leaf mass per area (LMA) is a key plant trait, reflecting tradeoffs between leaf photosynthetic function, longevity, and structural investment. Capturing spatial and temporal variability in LMA has been a long‐standing goal of ecological research and is an essential component for advancing Earth system models. Despite the substantial variation in LMA within and across Earth's biomes, an efficient, globally generalizable approach to predict LMA is still lacking.

    We explored the capacity to predict LMA from leaf spectra across much of the global LMA trait space, with values ranging from 17 to 393 g m−2. Our dataset contained leaves from a wide range of biomes from the high Arctic to the tropics, included broad‐ and needleleaf species, and upper‐ and lower‐canopy (i.e. sun and shade) growth environments.

    Here we demonstrate the capacity to rapidly estimate LMA using only spectral measurements across a wide range of species, leaf age and canopy position from diverse biomes. Our model captures LMA variability with high accuracy and low error (R2 = 0.89; root mean square error (RMSE) = 15.45 g m−2).

    Our finding highlights the fact that the leaf economics spectrum is mirrored by the leaf optical spectrum, paving the way for this technology to predict the diversity of LMA in ecosystems across global biomes.

     
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  6. Abstract

    Stomatal response to environmental conditions forms the backbone of all ecosystem and carbon cycle models, but is largely based on empirical relationships. Evolutionary theories of stomatal behaviour are critical for guarding against prediction errors of empirical models under future climates. Longstanding theory holds that stomata maximise fitness by acting to maintain constant marginal water use efficiency over a given time horizon, but a recent evolutionary theory proposes that stomata instead maximise carbon gain minus carbon costs/risk of hydraulic damage. Using data from 34 species that span global forest biomes, we find that the recent carbon‐maximisation optimisation theory is widely supported, revealing that the evolution of stomatal regulation has not been primarily driven by attainment of constant marginal water use efficiency. Optimal control of stomata to manage hydraulic risk is likely to have significant consequences for ecosystem fluxes during drought, which is critical given projected intensification of the global hydrological cycle.

     
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  7. Summary

    Deep‐water access is arguably the most effective, but under‐studied, mechanism that plants employ to survive during drought. Vulnerability to embolism and hydraulic safety margins can predict mortality risk at given levels of dehydration, but deep‐water access may delay plant dehydration. Here, we tested the role of deep‐water access in enabling survival within a diverse tropical forest community in Panama using a novel data‐model approach.

    We inversely estimated the effective rooting depth (ERD, as the average depth of water extraction), for 29 canopy species by linking diameter growth dynamics (1990–2015) to vapor pressure deficit, water potentials in the whole‐soil column, and leaf hydraulic vulnerability curves. We validated ERD estimates against existing isotopic data of potential water‐access depths.

    Across species, deeper ERD was associated with higher maximum stem hydraulic conductivity, greater vulnerability to xylem embolism, narrower safety margins, and lower mortality rates during extreme droughts over 35 years (1981–2015) among evergreen species. Species exposure to water stress declined with deeper ERD indicating that trees compensate for water stress‐related mortality risk through deep‐water access.

    The role of deep‐water access in mitigating mortality of hydraulically‐vulnerable trees has important implications for our predictive understanding of forest dynamics under current and future climates.

     
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