The process of sapwood/heartwood transition in trees is not fully understood. We tested whether the ontogenetically-stable apex-to-base conduit widening generates path length effects limiting the conductance of inner sapwood rings. The axial scaling ( b) of conduit hydraulic diameter ( Dh) was estimated at annual resolution in a spruce and beech tree. We compiled a global dataset of sapwood ring number ( NSWr), their average width ( SWrw), tree height ( H) and stem elongation rate ( ΔH) in conifer and angiosperm trees. A numerical model simulated the effects of H and ΔH on the conductance of each xylem ring ( KRING). b resulted ontogenetically stable. Simulations well predicted the observed patterns of increasing NSWr with H and decreasing NSWr with ΔH, assuming that heartwood forms when the marginal conductance gain of maintaining the functionality of an inner ring becomes negligible. Sapwood/heartwood transition minimizes the C costs associated to allocation to secondary growth and maintenance of living sapwood required to attain a given sapwood conductance. The number of sapwood rings depends on the effects of H and ΔH on the conductance of inner sapwood rings. The width of sapwood rings contributes to compensate for the lower conductance of inner sapwood rings at high ΔH.

The relationships between water potential and water content in plants and soil have long been of interest, and there is increasing focus on understanding how these fundamental measures are linked at larger spatial and temporal scales. In this Perspective, we explore how the theory of pressure-volume (PV) relationships can be applied at ecosystem scale. We define and evaluate the concept and limitations of the ecosystem and vegetation pressure-volume curves and discuss its application using existing data. As a proof of concept, plot-scale aboveground vegetation PV curves were generated from equilibrium (e.g. predawn) water potentials and water content of the above ground biomass of nine plots including tropical rainforest, savanna, temperate forest, and a long-term Amazonian rainforest drought experiment. Initial findings suggest high consistency among sites where the steady-state water:biomass ratio is approximately 1:3, while the relative values of ecosystem hydraulic capacitance and accessible water storage (the water volume between saturation and a threshold) do not vary systematically with biomass. The ecosystem-scale PV relationship provides a thermodynamically consistent steady-state view of ecosystem form and function and a biophysically robust basis for the interpretation of remote sensing data of vegetation and soil water content, with promise for revealing useful trends across ecosystems.

Transpiration is a key process driving energy, water and thus carbon dynamics. Global T products are fundamental for understanding and predicting vegetation processes. However, validation of these transpiration products is limited, mainly due to lack of suitable datasets. We propose a method to use SAPFLUXNET, the first quality-controlled global tree sap flow database, for evaluating transpiration products at global scale. Our method is based on evaluating temporal mismatches, rather than absolute values, by standardizing both transpiration and sap flow products. We evaluate how transpiration responses to hydro-meteorological variation from the Global Land Evaporation Amsterdam Model (GLEAM), a widely used global transpiration product, compare to in-situ responses from SAPFLUXNET field data. Our results show GLEAM and SAPFLUXNET temporal trends are in good agreement, but diverge under extreme conditions. Their temporal mismatches differ depending on the magnitude of transpiration and are not random, but linked to energy and water availability. Despite limitations, we show that the new global SAPFLUXNET dataset is a valuable tool to evaluate T products and identify problematic assumptions and processes embedded in models. The approach we propose can, therefore, be the foundation for a wider use of SAPFLUXNET, a new, independent, source of information, to understand the mechanisms controlling global transpiration fluxes.

We aim to identify the relative importance of vapour pressure deficit (VPD), soil water content (SWC) and photosynthetic photon flux density (PPFD) as drivers of tree canopy conductance, which is a key source of uncertainty for modelling vegetation responses under climate change. We use sap flow time series of 1858 trees in 122 sites from the SAPFLUXNET global database to obtain whole-tree canopy conductance (G). The coupling, defined as the percentage of variance (R2) of G explained by the three main hydrometeorological drivers (VPD, SWC and PPFD), was evaluated using linear mixed models. For each hydrometeorological driver we assess differences in coupling among biomes, and use multiple linear regression to explain R2 by climate, soil and vegetation structure. We found that in most areas tree canopy conductance is better explained by VPD than by SWC or PPFD. We also found that sites in drylands are less coupled to all three hydrometeorological drivers than those in other biomes. Climate, soil and vegetation structure were common controls of all three hydrometeorological couplings with G, with wetter climates, fine textured soils and tall vegetation being associated to tighter coupling. Differences across sites in the hydrometeorological coupling of tree canopy conductance may affect predictions of ecosystem dynamics under future climates, and should be accounted for explicitly in models.

It is easy to measure annual growth of a tree stem. It is hard to measure its daily growth. The reason for this difficulty is the microscopic scale and the need to separate processes that simultaneously result in reversible and irreversible stem expansion. Here we present a model that separates reversible from irreversible cell expansion. Our model is novel, because it explains reversible expansion as consequence of longitudinally and, importantly, radially transmitted changes of hydraulic and osmotic pressures in xylem and bark. To capture and quantify these changes, we manipulated daily stem growth by applying a phloem girdle to stems of 9-m tall trees. The model was informed by measurements of radial movement in stem tissues and sap flow before and after and positions below and above the girdle. Additional measurements of whole-crown fluxes of H2O and CO2, leaf water potentials, non-structural carbohydrates and respiration were used to document the physiological impacts of girdling. This work sheds new light on the role of radial transport processes underpinning daily growth of tree stems. The model helps explain diel patterns of stem growth in trees.

The future of tropical forests is dependent on the capacity of young trees to adjust to drought. We evaluated multiple hydraulic traits indicative of the drought tolerance of small trees across nine common genera at the world’s longest-running tropical throughfall exclusion experiment and compared their responses with surviving large canopy trees. Small understorey trees increased specific hydraulic conductivity by 56.3% and leaf:sapwood area ratio by 45.6% in response to the drought treatment. However, understorey trees in both a control and the throughfall exclusion treatment had significantly lower minimum stomatal conductance and maximum hydraulic leaf-specific conductivity relative to the large trees, as well as significantly greater hydraulic safety margin (HSM) and PLC and embolism resitance, occupying a distinctly different hydrualic niche. The greater HSM of small understorey trees relative to large canopy trees likely enables them to adjust other aspects of their hydraulic systems to take advantage of increases in light availability in the understorey, driven by drought-induced mortality of canopy trees. Our results suggest that small understorey trees can adjust their hydraulic systems in response to changes in water and light availability and this has major implications for the regeneration potential of tropical forests following droughts.

Hydraulic properties control plant responses to climate and are likely to be under strong selective pressure, but their macro-evolutionary history remains poorly characterized. To fill this gap, we compiled a global dataset of hydraulic traits describing xylem conductivity (Ks), xylem resistance to embolism (P50), sapwood allocation relative to leaf area (Hv) and drought exposure (ψmin), and matched it with global seed plant phylogenies. Individually, these traits present medium to high levels of phylogenetic signal, partly related to environmental selective pressures shaping lineage evolution. Most of these traits evolved independently of each other, being co-selected by the same environmental pressures. However, the evolutionary correlations between P50 and ψmin and between Ks and Hv show signs of deeper evolutionary integration because of functional, developmental or genetic constraints, conforming to evolutionary modules. We do not detect evolutionary integration between conductivity and resistance to embolism, rejecting a hardwired trade-off for this pair of traits.