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Abstract The cause of reduced leaf-level transpiration under elevated CO2 remains largely elusive. Here, we assessed stomatal, hydraulic, and morphological adjustments in a long-term experiment on Aleppo pine (Pinus halepensis) seedlings germinated and grown for 22–40 months under elevated (eCO2; c. 860 ppm) or ambient (aCO2; c. 410 ppm) CO2. We assessed if eCO2-triggered reductions in canopy conductance (gc) alter the response to soil or atmospheric drought and are reversible or lasting due to anatomical adjustments by exposing eCO2 seedlings to decreasing [CO2]. To quantify underlying mechanisms, we analyzed leaf abscisic acid (ABA) level, stomatal and leaf morphology, xylem structure, hydraulic efficiency, and hydraulic safety. Effects of eCO2 manifested in a strong reduction in leaf-level gc (−55%) not caused by ABA and not reversible under low CO2 (c. 200 ppm). Stomatal development and size were unchanged, while stomatal density increased (+18%). An increased vein-to-epidermis distance (+65%) suggested a larger leaf resistance to water flow. This was supported by anatomical adjustments of branch xylem having smaller conduits (−8%) and lower conduit lumen fraction (−11%), which resulted in a lower specific conductivity (−19%) and leaf-specific conductivity (−34%). These adaptations to CO2 did not change stomatal sensitivity to soil or atmospheric drought, consistent with similar xylem safety thresholds. In summary, we found reductions of gc under elevated CO2 to be reflected in anatomical adjustments and decreases in hydraulic conductivity. As these water savings were largely annulled by increases in leaf biomass, we do not expect alleviation of drought stress in a high CO2 atmosphere. 

Summary Embolism spreading in angiosperm xylem occurs via mesoporous pit membranes between vessels. Here, we investigate how the size of pore constrictions in pit membranes is related to pit membrane thickness and embolism resistance.Pit membranes were modelled as multiple layers to investigate how pit membrane thickness and the number of intervessel pits per vessel determine pore constriction sizes, the probability of encountering large pores, and embolism resistance. These estimations were complemented by measurements of pit membrane thickness, embolism resistance, and number of intervessel pits per vessel in stem xylem (n = 31, 31 and 20 species, respectively).The modelled constriction sizes in pit membranes decreased with increasing membrane thickness, explaining the measured relationship between pit membrane thickness and embolism resistance. The number of pits per vessel affected constriction size and embolism resistance much less than pit membrane thickness. Moreover, a strong relationship between modelled and measured embolism resistance was observed.Pore constrictions provide a mechanistic explanation for why pit membrane thickness determines embolism resistance, which suggests that hydraulic safety can be uncoupled from hydraulic efficiency. Although embolism spreading remains puzzling and encompasses more than pore constriction sizes, angiosperms are unlikely to have leaky pit membranes, which enables tensile transport of water. 

Abstract. Plant transpiration links physiological responses ofvegetation to water supply and demand with hydrological, energy, and carbonbudgets at the land–atmosphere interface. However, despite being the mainland evaporative flux at the global scale, transpiration and its response toenvironmental drivers are currently not well constrained by observations.Here we introduce the first global compilation of whole-plant transpirationdata from sap flow measurements (SAPFLUXNET, https://sapfluxnet.creaf.cat/, last access: 8 June 2021).We harmonized and quality-controlled individual datasets supplied bycontributors worldwide in a semi-automatic data workflow implemented in theR programming language. Datasets include sub-daily time series of sap flowand hydrometeorological drivers for one or more growing seasons, as well asmetadata on the stand characteristics, plant attributes, and technicaldetails of the measurements. SAPFLUXNET contains 202 globally distributeddatasets with sap flow time series for 2714 plants, mostly trees, of 174species. SAPFLUXNET has a broad bioclimatic coverage, withwoodland/shrubland and temperate forest biomes especially well represented(80 % of the datasets). The measurements cover a wide variety of standstructural characteristics and plant sizes. The datasets encompass theperiod between 1995 and 2018, with 50 % of the datasets being at least 3 years long. Accompanying radiation and vapour pressure deficit data areavailable for most of the datasets, while on-site soil water content isavailable for 56 % of the datasets. Many datasets contain data for speciesthat make up 90 % or more of the total stand basal area, allowing theestimation of stand transpiration in diverse ecological settings. SAPFLUXNETadds to existing plant trait datasets, ecosystem flux networks, and remotesensing products to help increase our understanding of plant water use,plant responses to drought, and ecohydrological processes. SAPFLUXNET version0.1.5 is freely available from the Zenodo repository (https://doi.org/10.5281/zenodo.3971689; Poyatos et al., 2020a). The“sapfluxnetr” R package – designed to access, visualize, and processSAPFLUXNET data – is available from CRAN.