Forests out of balance: The impacts of drought-induced tree mortality on carbon and water cycling
Forestry
Final Report Abstract
Trees and forests are affected by a rapidly changing climate. The rise in frequency and severity of extreme drought events, exemplified by the conditions in Germany during the exceptionally dry years of 2018 and 2019, has led to reduced tree vitality and increased mortality. Despite the common assumption in the drought tolerance of pine trees, Scots pine experienced widespread mortality during these extreme drought years. However, whether this trend will persist, diminish, or escalate remains uncertain, primarily due to gaps in our understanding of drought resilience processes, especially concerning how stress impacts impede post-stress recovery. This project concentrated on investigating the responses of pine trees to drought and heat stress, as well as their capacity to recover from such stress events, by integrating experimental and modeling approaches. Our experimental findings emphasized the tight regulation of the carbon (C) balance in pine trees: a reduction in C gain from photosynthesis led to corresponding decreases in respiration and growth. During periods of water scarcity and low assimilation, both above- and belowground growth ceased abruptly, accompanied by a decline in starch reserves. This suggests that pine trees can swiftly respond to stress and maintain a relatively balanced C metabolism, thus avoiding C starvation. However, as drought persists, dehydration and damage to the hydraulic system intensify. Furthermore, we demonstrated that the impacts of drought stress exacerbate with increased atmospheric dryness and heat stress, and elevated CO2 levels do not ameliorate the stress response. Upon release from severe drought, Scots pine seedlings were unable to repair damage to the water transport system, delaying the recovery of photosynthesis and growth, while carbohydrate reserves accumulated. In addition to these adverse responses, we observed that stress can prompt structural adjustments, such as reductions in leaf area, which can decrease tree water loss and mitigate drought stress in the longer-term. To evaluate these processes at the forest level, we incorporated a detailed tree hydraulic description into an advanced ecosystem model. The model accurately depicted the impacts of drought by simulating a reduction in hydraulic conductance, leading to losses in leaf area and sapwood area. We successfully applied the model to a semi-arid pine site, which is welladapted to seasonal drought conditions. Our model findings aligned with observations, showing minimal loss of hydraulic functioning in the semi-arid forest. This contrasts with the severe drought responses observed in temperate forests following the 2018/2019 summer droughts, which were captured by the model through damages of leaf and sapwood area. Albeit often overlooked by models, simulating these extreme stress damages is crucial for evaluating the long-term effects of climate change on forest ecosystems.
Publications
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Research frontiers for improving our understanding of drought‐induced tree and forest mortality. New Phytologist, 218(1), 15-28.
Hartmann, Henrik; Moura, Catarina F.; Anderegg, William R. L.; Ruehr, Nadine K.; Salmon, Yann; Allen, Craig D.; Arndt, Stefan K.; Breshears, David D.; Davi, Hendrik; Galbraith, David; Ruthrof, Katinka X.; Wunder, Jan; Adams, Henry D.; Bloemen, Jasper; Cailleret, Maxime; Cobb, Richard; Gessler, Arthur; Grams, Thorsten E. E.; Jansen, Steven ... & O'Brien, Michael
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Beyond the extreme: recovery of carbon and water relations in woody plants following heat and drought stress. Tree Physiology, 39(8), 1285-1299.
Ruehr, Nadine K.; Grote, Rüdiger; Mayr, Stefan & Arneth, Almut
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Climate and plant trait strategies determine tree carbon allocation to leaves and mediate future forest productivity. Global Change Biology, 25(10), 3395-3405.
Trugman, Anna T.; Anderegg, Leander D. L.; Wolfe, Brett T.; Birami, Benjamin; Ruehr, Nadine K.; Detto, Matteo; Bartlett, Megan K. & Anderegg, William R. L.
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One Century of Forest Monitoring Data in Switzerland Reveals Species- and Site-Specific Trends of Climate-Induced Tree Mortality. Frontiers in Plant Science, 10.
Etzold, Sophia; Ziemińska, Kasia; Rohner, Brigitte; Bottero, Alessandra; Bose, Arun K.; Ruehr, Nadine K.; Zingg, Andreas & Rigling, Andreas
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A first assessment of the impact of the extreme 2018 summer drought on Central European forests. Basic and Applied Ecology, 45, 86-103.
Schuldt, Bernhard; Buras, Allan; Arend, Matthias; Vitasse, Yann; Beierkuhnlein, Carl; Damm, Alexander; Gharun, Mana; Grams, Thorsten E.E.; Hauck, Markus; Hajek, Peter; Hartmann, Henrik; Hiltbrunner, Erika; Hoch, Günter; Holloway-Phillips, Meisha; Körner, Christian; Larysch, Elena; Lübbe, Torben; Nelson, Daniel B.; Rammig, Anja ... & Kahmen, Ansgar
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Drought-Induced Xylem Embolism Limits the Recovery of Leaf Gas Exchange in Scots Pine. Plant Physiology, 184(2), 852-864.
Rehschuh, Romy; Cecilia, Angelica; Zuber, Marcus; Faragó, Tomáš; Baumbach, Tilo; Hartmann, Henrik; Jansen, Steven; Mayr, Stefan & Ruehr, Nadine
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Dying by drying: Timing of physiological stress thresholds related to tree death is not significantly altered by highly elevated CO2. Plant, Cell & Environment, 44(2), 356-370.
Gattmann, Marielle; Birami, Benjamin; Nadal, Sala Daniel & Ruehr, Nadine Katrin
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Hot drought reduces the effects of elevated CO2 on tree water‐use efficiency and carbon metabolism. New Phytologist, 226(6), 1607-1621.
Birami, Benjamin; Nägele, Thomas; Gattmann, Marielle; Preisler, Yakir; Gast, Andreas; Arneth, Almut & Ruehr, Nadine K.
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Assessing model performance via the most limiting environmental driver in two differently stressed pine stands. Ecological Applications, 31(4).
Nadal‐Sala, Daniel; Grote, Rüdiger; Birami, Benjamin; Lintunen, Anna; Mammarella, Ivan; Preisler, Yakir; Rotenberg, Eyal; Salmon, Yann; Tatarinov, Fedor; Yakir, Dan & Ruehr, Nadine K.
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Diverging responses of water and carbon relations during and after heat and hot drought stress in Pinus sylvestris. Tree Physiology, 42(8), 1532-1548.
Rehschuh, Romy & Ruehr, Nadine K.
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Heatwave frequency and seedling death alter stress-specific emissions of volatile organic compounds in Aleppo pine. Oecologia, 197(4), 939-956.
Birami, Benjamin; Bamberger, Ines; Ghirardo, Andrea; Grote, Rüdiger; Arneth, Almut; Gaona-Colmán, Elizabeth; Nadal-Sala, Daniel & Ruehr, Nadine K.
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Leaf Shedding and Non-Stomatal Limitations of Photosynthesis Mitigate Hydraulic Conductance Losses in Scots Pine Saplings During Severe Drought Stress. Frontiers in Plant Science, 12.
Nadal-Sala, Daniel; Grote, Rüdiger; Birami, Benjamin; Knüver, Timo; Rehschuh, Romy; Schwarz, Selina & Ruehr, Nadine K.
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Tree allocation dynamics beyond heat and hot drought stress reveal changes in carbon storage, belowground translocation and growth. New Phytologist, 233(2), 687-704.
Rehschuh, Romy; Rehschuh, Stephanie; Gast, Andreas; Jakab, Andrea‐Livia; Lehmann, Marco M.; Saurer, Matthias; Gessler, Arthur & Ruehr, Nadine K.
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Anatomical adjustments of the tree hydraulic pathway decrease canopy conductance under long-term elevated CO2. Plant Physiology, 191(1), 252-264.
Gattmann, Marielle; McAdam, Scott A. M.; Birami, Benjamin; Link, Roman; Nadal-Sala, Daniel; Schuldt, Bernhard; Yakir, Dan & Ruehr, Nadine K.
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Forest canopy mortality during the 2018-2020 summer drought years in Central Europe: The application of a deep learning approach on aerial images across Luxembourg. Forestry: An International Journal of Forest Research, 97(3), 376-387.
Schwarz, Selina; Werner, Christian; Fassnacht, Fabian Ewald & Ruehr, Nadine K.
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Integration of tree hydraulic processes and functional impairment to capture the drought resilience of a semi-arid pine forest.
Nadal-Sala, Daniel; Grote, Rüdiger; Kraus, David; Hochberg, Uri; Klein, Tamir; Wagner, Yael; Tatarinov, Fedor; Yakir, Dan & Ruehr, Nadine Katrin
