Forest ecosystems play a central role in global water and carbon cycles, yet the impact of global climate change, in particular drought, on trees and forests is poorly understood. Therefore, there is an urgent need for forest-scale experiments in improving our understanding of trees' responses to extreme drought events and subsequent recovery under field conditions. Here, we present the design and efficacy of a novel throughfall exclusion experiment with retractable roofs in a mature forest allowing for flexible drought and recovery periods. A total of 12 plots (144 AE 26 m 2 on average) with 3-7 European beech and Norway spruce trees each were established by root trenching to a depth of one meter, four years prior to the experiment. Subsequent installation of roofs (n = 6) allowed for the removal of throughfall precipitation and almost a complete non-availability of soil water in the upper 70 cm during five subsequent growing seasons, that is, 2014-2018. This reduction in available soil water resulted in pre-dawn leaf water potentials down to −1.8 MPa in mature trees. Stem diameter growth decreased by 30% in beech and 70% in spruce, and fine root abundance was reduced by 57% in beech and 73% in spruce compared with controls. After only one growing season, the mycorrhizal community composition changed in response to drought. Careful watering of hydrophobic forest soils in early summer of 2019 resulted in recovered pre-dawn leaf water potentials of drought-stressed trees within one week. Recovery of stem diameter growth, however, did not occur within the same growing season and remained reduced by 33% in beech and 69% in spruce compared with controls. The implemented throughfall exclusion system imposed recurrent seasonal drought events on a mature beech/spruce forest with high efficacy. Shifts in community composition of mycorrhizae in parallel to tree growth decline advocate for a more holistic view on forest-scale drought and watering experiments, particularly in light of more frequently predicted drought events in future. The perennial nature of mature trees and their subsequent slow recovery from drought, that is, over multiple growing seasons, argues for more long-term experiments that span several years.

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Hydraulic failure and carbon starvation are recognized as main causes of drought-induced forest decline. As water transport and carbon dynamics are strictly interdependent, it is necessary to clarify how dehydration-rehydration cycles are affecting the relations between stem embolism and non-structural carbohydrates (NSC). This is particularly needed for conifers whose embolism repair capability is still controversial. Potted Norway spruce saplings underwent two drought-re-irrigation cycles of same intensity, but performed in two consecutive summers. During the second cycle, stem percent loss of hydraulic conductivity (PLC) and NSC content showed no carry-over effects from the previous drought, indicating complete long-term recovery. The second drought treatment induced moderate PLC (20%) and did not affect total NSCs content, while starch was converted to soluble sugars in the bark. After one week of re-irrigation, PLC recovered to pre-stress values (0%) and NSCs were depleted, only in the wood, by about 30%. Our data suggest that spruce can repair xylem embolism and that, when water is newly available, NSCs stored in xylem parenchyma can be mobilized over short term to sustain respiration and/or for processes involved in xylem transport restoration. This, however, might imply dependency on sapwood NSC reserves for survival, especially if frequent drought spells occur.

Decreasing water availability due to climate change poses the question of whether and to what extent tree species are able to hydraulically acclimate and how hydraulic traits of stems and leaves are coordinated under drought. In a through-fall exclusion experiment, hydraulic acclimation was analyzed in a mixed forest stand of Fagus sylvatica L. and Picea abies (L.) Karst. In drought-stressed (TE, through-fall exclusion over 2 years) and control (CO) trees, hydraulic vulnerability was studied in branches as well as in leaves (F. sylvatica) and end-twigs (P. abies, entirely formed during the drought period) sampled at the same height in sun-exposed portions of the tree crown. In addition, relevant xylem anatomical traits and leaf pressure-volume relations were analyzed. The TE trees reached pre-dawn water potentials down to -1.6 MPa. In both species, water potentials at 50% loss of xylem hydraulic conductivity were ~0.4 MPa more negative in TE than in CO branches. Foliage hydraulic vulnerability (expressed as water potential at 50% loss of leaf/end-twig hydraulic conductance) and water potential at turgor loss point were also, respectively, 0.4 and 0.5 MPa lower in TE trees. Minor differences were observed in conduit mean hydraulic diameter and cell wall reinforcement. Our findings indicate significant and fast hydraulic acclimation under relatively mild drought in both tree species. Acclimation was well coordinated between branches and foliage, which might be essential for survival and productivity of mature trees under future drought periods.

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