Balancing the risks of hydraulic failure and carbon starvation : a twig scale analysis in declining Scots pine

Understanding physiological processes involved in drought-induced mortality is important for predicting the future of forests and for modelling the carbon and water cycles. Recent research has highlighted the variable risks of carbon starvation and hydraulic failure in drought-exposed trees. However, little is known about the specific responses of leaves and supporting twigs, despite their critical role in balancing carbon acquisition and water loss. Comparing healthy (non-defoliated) and unhealthy (defoliated) Scots pine at the same site, we measured the physiological variables involved in regulating carbon and water resources. Defoliated trees showed different responses to summer drought compared with non-defoliated trees. Defoliated trees maintained gas exchange while non-defoliated trees reduced photosynthesis and transpiration during the drought period. At the branch scale, very few differences were observed in non-structural carbohydrate concentrations between health classes. However, defoliated trees tended to have lower water potentials and smaller hydraulic safety margins. While non-defoliated trees showed a typical response to drought for an isohydric species, the physiology appears to be driven in defoliated trees by the need to maintain carbon resources in twigs. These responses put defoliated trees at higher risk of branch hydraulic failure and help explain the interaction between carbon starvation and hydraulic failure in dying trees.Peer reviewe

How woody plants adjust above- and below-ground traits in response to sustained drought

This is the final version. Available on open access from Wiley via the DOI in this recordFuture increases in drought severity and frequency are predicted to have substantial impacts on plant function and survival. However, there is considerable uncertainty concerning what drought adjustment is and whether plants can adjust to sustained drought. This review focuses on woody plants and synthesises the evidence for drought adjustment in a selection of key above-ground and below-ground plant traits. We assess whether evaluating the drought adjustment of single traits, or selections of traits that operate on the same plant functional axis (e.g. photosynthetic traits) is sufficient, or whether a multi-trait approach, integrating across multiple axes, is required. We conclude that studies on drought adjustments in woody plants might overestimate the capacity for adjustment to drier environments if spatial studies along gradients are used, without complementary experimental approaches. We provide evidence that drought adjustment is common in above-ground and below-ground traits; however, whether this is adaptive and sufficient to respond to future droughts remains uncertain for most species. To address this uncertainty, we must move towards studying trait integration within and across multiple axes of plant function (e.g. above-ground and below-ground) to gain a holistic view of drought adjustments at the whole-plant scale and how these influence plant survival.Natural Environment Research Council (NERC)MINECOEuropean Union Horizon 202

Transpiration from subarctic deciduous woodlands: environmental controls and contribution to ecosystem evapotranspiration

Potential land‐climate feedbacks in subarctic regions, where rapid warming is driving forest expansion into the tundra, may be mediated by differences in transpiration of different plant functional types. Here we assess the environmental controls of overstorey transpiration and its relevance for ecosystem evapotranspiration in subarctic deciduous woodlands. We measured overstorey transpiration of mountain birch canopies and ecosystem evapotranspiration in two locations in northern Fennoscandia, having dense (Abisko) and sparse (Kevo) overstories. For Kevo, we also upscale chamber‐measured understorey evapotranspiration from shrubs and lichen using a detailed land cover map. Sub‐daily evaporative fluxes were not affected by soil moisture, and showed similar controls by vapour pressure deficit and radiation across sites. At the daily timescale, increases in evaporative demand led to proportionally higher contributions of overstorey transpiration to ecosystem evapotranspiration. For the entire growing season, the overstorey transpired 33% of ecosystem evapotranspiration in Abisko and only 16% in Kevo. At this latter site, the understorey had a higher leaf area index and contributed more to ecosystem evapotranspiration compared to the overstorey birch canopy. In Abisko, growing season evapotranspiration was 27% higher than precipitation, consistent with a gradual soil moisture depletion over the summer. Our results show that overstorey canopy transpiration in subarctic deciduous woodlands is not the dominant evaporative flux. However, given the observed environmental sensitivity of evapotranspiration components, the role of deciduous trees in driving ecosystem evapotranspiration may increase with the predicted increases in tree cover and evaporative demand across subarctic regions

Bridging scales: An approach to evaluate the temporal patterns of global transpiration products using tree‐scale sap flow data

This is the final version. Available from Wiley via the DOI in this record.Data Availability Statement All data used in this work is freely available at the GLEAM (https://gleam.io/) and SAPFLUXNET (http://sapfluxnet.creaf.cat/) online repositoriesTranspiration is a key process driving energy, water and thus carbon dynamics. Global transpiration products are fundamental for understanding and predicting vegetation processes. However, validation of these transpiration products is limited, mainly due to lack of suitable data sets. We propose a method to use SAPFLUXNET, the first quality-controlled global tree sap flow (SF) 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 SF 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 data set 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.Natural Environment Research Council (NERC)Natural Environment Research Council (NERC)Ministerio de Ciencia e InnovaciónEuropean Research CouncilNewton FundHumboldt Fellowship for Experienced Researcher

The Impact of a Simple Representation of Non-Structural Carbohydrates on the Simulated Response of Tropical Forests to Drought

This is the final version. Available on open access from European Geosciences Union via the DOI in this recordCode availability. A model example of SUGAR for a single site and set-up to run at Caxiuanã using output from JULES is available at https://doi.org/10.5281/zenodo.3547613 (Jones, 2019). For further information or code please contact [email protected] representing the response of ecosystems to environmental change in land surface models (LSMs) is crucial to making accurate predictions of future climate. Many LSMs do not correctly capture plant respiration and growth fluxes, particularly in response to extreme climatic events. This is in part due to the unrealistic assumption that total plant carbon expenditure (PCE) is always equal to gross carbon accumulation by photosynthesis. We present and evaluate a simple model of labile carbon storage and utilisation (SUGAR) designed to be integrated into an LSM, which allows simulated plant respiration and growth to vary independent of photosynthesis. SUGAR buffers simulated PCE against seasonal variation in photosynthesis, producing more constant (less variable) predictions of plant growth and respiration relative to an LSM that does not represent labile carbon storage. This allows the model to more accurately capture observed carbon fluxes at a large-scale drought experiment in a tropical moist forest in the Amazon, relative to the Joint UK Land Environment Simulator LSM (JULES). SUGAR is designed to improve the representation of carbon storage in LSMs and provides a simple framework that allows new processes to be integrated as the empirical understanding of carbon storage in plants improves. The study highlights the need for future research into carbon storage and allocation in plants, particularly in response to extreme climate events such as drought.Natural Environment Research Council (NERC)Newton FundAustralian Research Council (ARC)Spanish Ministry of Economy and Competitiveness (MINECO)Engineering and Physical Sciences Research Council (EPSRC)JPL-Caltech President's and Director's Research & Development FundMet Office Hadley Centre Climate ProgrammeEuropean Union Horizon 202

Stand dynamics modulate water cycling and mortality risk in droughted tropical forest

This is the author accepted manuscript. The final version is available from Wiley via the DOI in this record.Transpiration from the Amazon rainforest generates an essential water source at a global and local scale. However, changes in rainforest function with climate change can disrupt this process, causing significant reductions in precipitation across Amazonia, and potentially at a global scale. We report the only study of forest transpiration following a long-term (>10 year) experimental drought treatment in Amazonian forest. After 15 years of receiving half the normal rainfall, drought-related tree mortality caused total forest transpiration to decrease by 30%. However, the surviving droughted trees maintained or increased transpiration because of reduced competition for water and increased light availability, which is consistent with increased growth rates. Consequently, the amount of water supplied as rainfall reaching the soil and directly recycled as transpiration increased to 100%. This value was 25% greater than for adjacent nondroughted forest. If these drought conditions were accompanied by a modest increase in temperature (e.g., 1.5°C), water demand would exceed supply, making the forest more prone to increased tree mortality.This work is a product of UK NERC grant NE/J011002/1 to PM and MM, CNPQ grant 457914/2013-0/MCTI/CNPq/FNDCT/LBA/ESECAFLOR to ACLD, an ARC grant FT110100457 to PM and a UK NERC independent fellowship grant NE/N014022/1 to LR. It was previously supported by NERC NER/A/S/2002/00487, NERC GR3/11706, EU FP5-Carbonsink and EU FP7-Amazalert to PM. RP acknowledges support of MINECO (Spain), grant CGL2014-5583-JIN