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Sack, Lawren

Publications and source records attributed to Sack, Lawren.

The ecosystem wilting point defines drought response and recovery of a Quercus‐Carya forest

Abstract Soil and atmospheric droughts increasingly threaten plant survival and productivity around the world. Yet, conceptual gaps constrain our ability to predict ecosystem‐scale drought impacts under climate change. Here, we introduce the ecosystem wilting point (Ψ EWP ), a property that integrates the drought response of an ecosystem's plant community across the soil–plant–atmosphere continuum. Specifically, Ψ EWP defines a threshold below which the capacity of the root system to extract soil water and the ability of the leaves to maintain stomatal function are strongly diminished. We combined ecosystem flux and leaf water potential measurements to derive the Ψ EWP of a Quercus‐Carya forest from an “ecosystem pressure–volume (PV) curve,” which is analogous to the tissue‐level technique. When community predawn leaf water potential (Ψ pd ) was above Ψ EWP (=−2.0 MPa), the forest was highly responsive to environmental dynamics. When Ψ pd fell below Ψ EWP , the forest became insensitive to environmental variation and was a net source of carbon dioxide for nearly 2 months. Thus, Ψ EWP is a threshold defining marked shifts in ecosystem functional state. Though there was rainfall‐induced recovery of ecosystem gas exchange following soaking rains, a legacy of structural and physiological damage inhibited canopy photosynthetic capacity. Although over 16 growing seasons, only 10% of Ψ pd observations fell below Ψ EWP , the forest is commonly only 2–4 weeks of intense drought away from reaching Ψ EWP , and thus highly reliant on frequent rainfall to replenish the soil water supply. We propose, based on a bottom‐up analysis of root density profiles and soil moisture characteristic curves, that soil water acquisition capacity is the major determinant of Ψ EWP , and species in an ecosystem require compatible leaf‐level traits such as turgor loss point so that leaf wilting is coordinated with the inability to extract further water from the soil.

54 ENVIRONMENTAL SCIENCES↗

Leaves as bottlenecks: The contribution of tree leaves to hydraulic resistance within the soil–plant–atmosphere continuum

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 (R leaf ) to total soil-to-leaf hydraulic resistance (R total ), or fR leaf (= R leaf /R total ), is thought to be large, but this has not been tested comprehensively. Here, we compiled a multi-biome dataset of fR leaf using new and previously published measurements of pressure differences within trees in situ. Across 80 samples, fR leaf averaged 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 fR leaf for 19 tree samples, which was 0.40 (95% CI = 0.29-0.56). The in-situ technique produces a more accurate descriptor of fR leaf because it accounts for dynamic leaf hydraulic conductance. Both approaches demonstrate the outsized role of leaves in controlling tree hydrodynamics. A larger fR leaf may play a role in protecting stems from loss of hydraulic conductance. Thus, the decline in fR leaf with tree height would contribute to greater drought vulnerability in taller trees and potentially to their observed disproportionate drought mortality.

54 ENVIRONMENTAL SCIENCES↗

Leaf habit affects the distribution of drought sensitivity but not water transport efficiency in the tropics

Abstract Considering the global intensification of aridity in tropical biomes due to climate change, we need to understand what shapes the distribution of drought sensitivity in tropical plants. We conducted a pantropical data synthesis representing 1117 species to test whether xylem‐specific hydraulic conductivity (K S ), water potential at leaf turgor loss (Ψ TLP ) and water potential at 50% loss of K S (Ψ P50 ) varied along climate gradients. The Ψ TLP and Ψ P50 increased with climatic moisture only for evergreen species, but K S did not. Species with high Ψ TLP and Ψ P50 values were associated with both dry and wet environments. However, drought‐deciduous species showed high Ψ TLP and Ψ P50 values regardless of water availability, whereas evergreen species only in wet environments. All three traits showed a weak phylogenetic signal and a short half‐life. These results suggest strong environmental controls on trait variance, which in turn is modulated by leaf habit along climatic moisture gradients in the tropics.

54 ENVIRONMENTAL SCIENCES↗

Leaf water potential measurements using the pressure chamber: Synthetic testing of assumptions towards best practices for precision and accuracy

Leaf water potential (ψ leaf ), typically measured using the pressure chamber, is the most important metric of plant water status, providing high theoretical value and information content for multiple applications in quantifying critical physiological processes including drought responses. Pressure chamber measurements of ψ leaf (ψ leafPC ) are most typical, yet, the practical complexity of the technique and of the underlying theory has led to ambiguous understanding of the conditions to optimize measurements. Consequently, specific techniques and precautions diversified across the global research community, raising questions of reliability and repeatability. Here, we surveyed specific methods of ψ leafPC from multiple laboratories, and synthesized experiments testing common assumptions and practices in ψ leafPC for diverse species: (i) the need for equilibration of previously transpiring leaves; (ii) leaf storage before measurement; (iii) the equilibration of ψ leaf for leaves on bagged branches of a range of dehydration; (iv) the equilibration of ψ leaf across the lamina for bagged leaves, and the accuracy of measuring leaves with artificially ‘elongated petioles’; (v) the need in ψ leaf measurements for bagging leaves and high humidity within the chamber; (vi) the need to avoid liquid water on leaf surfaces; (vii) the use of ‘pulse’ pressurization versus gradual pressurization; and (viii) variation among experimenters in ψ leafPC determination. Based on our findings we provide a best practice protocol to maximise accuracy, and provide recommendations for ongoing species-specific tests of important assumptions in future studies.

54 ENVIRONMENTAL SCIENCES↗

Detecting forest response to droughts with global observations of vegetation water content

Droughts in a warming climate have become more common and more extreme, making understanding forest responses to water stress increasingly pressing. Analysis of water stress in trees has long focused on water potential in xylem and leaves, which influences stomatal closure and water flow through the soil-plant-atmosphere continuum. At the same time, changes of vegetation water content (VWC) are linked to a range of tree responses, including fluxes of water and carbon, mortality, flammability, and more. Unlike water potential, which requires demanding in situ measurements, VWC can be retrieved from remote sensing measurements, particularly at microwave frequencies using radar and radiometry. Here, we highlight key frontiers through which VWC has the potential to significantly increase our understanding of forest responses to water stress. To validate remote sensing observations of VWC at landscape scale and to better relate them to data assimilation model parameters, we introduce an ecosystem-scale analogue of the pressure-volume curve, the non-linear relationship between 44 average leaf or branch water potential and water content commonly used in plant hydraulics. The 45 sources of variability in these ecosystem-scale pressure-volume curves and their relationship to 46 forest response to water stress are discussed. We further show to what extent diel, seasonal, and 47 decadal dynamics of VWC reflect variations in different processes relating the tree response to 48 water stress. VWC can also be used for inferring belowground conditions – which are difficult to 49 impossible to observe directly. Lastly, we discuss how a dedicated geostationary spaceborne 50 observational system for VWC, when combined with existing datasets, can capture diel and 51 seasonal water dynamics to advance the science and applications of global forest vulnerability to 52 future droughts.

54 ENVIRONMENTAL SCIENCES↗

Hydraulically‐vulnerable trees survive on deep‐water access during droughts in a tropical forest

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.

54 ENVIRONMENTAL SCIENCES↗

Tree Canopies Reflect Mycorrhizal Composition

Mycorrhizae alter global patterns of CO 2 fertilization, carbon storage, and elemental cycling, yet knowledge of their global distributions is currently limited by the availability of forest inventory data. Here, we show that maps of tree-mycorrhizal associations (hereafter “mycorrhizal maps”) can be improved by the novel technology of imaging spectroscopy because mycorrhizal signatures propagate up from plant roots to impact forest canopy chemistry. We analyzed measurements from 143 airborne imaging spectroscopy surveys over 112,975 individual trees collected across 13 years. Results show remarkable accuracy in capturing ground truth observations of mycorrhizal associations from canopy signals across disparate landscapes (R 2 = 0.92, p < 0.01). Additionally, upcoming imaging spectroscopy satellite missions can reveal new insights into landscape-scale variations in water, nitrogen, phosphorus, carotenoid/anthocyanin, and cellulose/lignin composition. Applied globally, this approach could improve the spatial precision of mycorrhizal distributions by a factor of roughly 104 and facilitate the incorporation of dynamic shifts in forest composition into Earth system models.

58 GEOSCIENCES↗

Leaf turgor loss point shapes local and regional distributions of evergreen but not deciduous tropical trees

Summary The effects of climate change on tropical forests will depend on how diverse tropical tree species respond to drought. Current distributions of evergreen and deciduous tree species across local and regional moisture gradients reflect their ability to tolerate drought stress, and might be explained by functional traits. We measured leaf water potential at turgor loss (i.e. ‘wilting point’; π tlp ), wood density (WD) and leaf mass per area (LMA) on 50 of the most abundant tree species in central Panama. We then tested their ability to explain distributions of evergreen and deciduous species within a 50 ha plot on Barro Colorado Island and across a 70 km rainfall gradient spanning the Isthmus of Panama. Among evergreen trees, species with lower π tlp were associated with drier habitats, with π tlp explaining 28% and 32% of habitat association on local and regional scales, respectively, greatly exceeding the predictive power of WD and LMA. In contrast, π tlp did not predict habitat associations among deciduous species. Across spatial scales, π tlp is a useful indicator of habitat preference for tropical tree species that retain their leaves during periods of water stress, and holds the potential to predict vegetation responses to climate change.

59 BASIC BIOLOGICAL SCIENCES↗

Soil Water Potentials (1990-2018) from a calibrated ELM-FATES, and rooting depth analyses scripts, PA-BCI, Panama

This dataset has ELM-FATES simulation outputs and scripts that accompany and reproduce the manuscript, Chitra-Tarak et al. (2021) Hydraulically-vulnerable trees survive on deep-water access during droughts in a tropical forest. New Phytologist. https: //doi.org/10.1111/nph.17464”. This dataset is organized into four parts: (1) source code for the model, Energy Exascale Earth System Land Model-Functionally Assembled Advanced Terrestrial simulator (ELM-FATES) (2) R scripts associated with ELM-FATES simulations: to prepare files for parameter ensemble simulation runs, calibrate the model with observed hydrological fluxes observed at BCI, re-run the model with tuned parameters and obtain long-term soil matric potentials by depth (3) ELM-FATES hydrological simulation outputs at PA-BCI, Panama (1990-2018, daily), and (4) R scripts for inverse rooting depth modeling and post-processing: prepare datasets and inversely model tree rooting depths at BCI, test hypotheses about the relationship of rooting depths with plant hydraulic traits and drought-induced mortality, and reproduce the manuscript itself. n.b. 1. Data needed to run R scripts in (2) and (4) and thus to reproduce the manuscript are sourced from various public datasets. Refer to the data availability section of the accompanying manuscript. 2. This dataset version (prepared on May 12, 2021) matches the revised and published version of the accompanying manuscript (published May 16, 2021) replacing a previous version. In the publication, this dataset is cited as follows: Chitra-Tarak R, Xu C, Aguilar S, Anderson-Teixeira K, Chambers J, Detto M, Faybishenko B, Fisher R, Knox R, Koven C et al. 2020. Soil water potentials (1990–2018) from a calibrated ELM-FATES, and rooting depth analyses scripts, PA-BCI, Panama. 2.0. NGEE Tropics Data Collection. doi: 10.15486/ngt/1696806

54 ENVIRONMENTAL SCIENCES↗

Global root traits (GRooT) database

Motivation: Trait data are fundamental to the quantitative description of plant form and function. Although root traits capture key dimensions related to plant responses to changing environmental conditions and effects on ecosystem processes, they have rarely been included in large-scale comparative studies and global models. For instance, root traits remain absent from nearly all studies that define the global spectrum of plant form and function. Thus, to overcome conceptual and methodological roadblocks preventing a widespread integration of root trait data into large-scale analyses we created the Global Root Trait (GRooT) Database. GRooT provides readyto- use data by combining the expertise of root ecologists with data mobilization and curation. Specifically, we (a) determined a set of core root traits relevant to the description of plant form and function based on an assessment by experts, (b) maximized species coverage through data standardization within and among traits, and (c) implemented data quality checks. Main types of variables contained: GRooT contains 114,222 trait records on 38 continuous root traits. Spatial location and grain: Global coverage with data from arid, continental, polar, temperate and tropical biomes. Data on root traits were derived from experimental studies and field studies. Time period and grain: Data were recorded between 1911 and 2019. Major taxa and level of measurement: GRooT includes root trait data for which taxonomic information is available. Trait records vary in their taxonomic resolution, with subspecies or varieties being the highest and genera the lowest taxonomic resolution available. It contains information for 184 subspecies or varieties, 6,214 species, 1,967 genera and 254 families. Owing to variation in data sources, trait records in the database include both individual observations and mean values. Software format: GRooT includes two csv files. A GitHub repository contains the csv files and a script in R to query the database.

59 BASIC BIOLOGICAL SCIENCES↗