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.
Hydraulic redistribution (HR) of soil water through plant roots is a crucial phenomenon improving the water balance of plants and ecosystems. It is mostly described under severe drought, and not yet studied under moderate drought. We tested the potential of HR under moderate drought, hypothesizing that (H1) tree species redistribute soil water in their roots even under moderate drought and that (H2) neighboring plants are supported with water provided by redistributing plants. Trees were planted in split-root systems with one individual (i.e., split-root plant, SRP) having its roots divided between two pots with one additional tree each. Species were 2- to 4-year-old English oak (Quercus robur L.), European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst). A gradient in soil water potential (ψsoil) was established between the two pots (-0.55 ± 0.02 MPa and -0.29 ± 0.03 MPa), and HR was observed by labeling with deuterium-enriched water. Irrespective of species identity, 93% of the SRPs redistributed deuterium enriched water from the moist to the drier side, supporting H1. Eighty-eight percent of the plants in the drier pots were deuterium enriched in their roots, with 61 ± 6% of the root water originating from SRP roots. Differences in HR among species were related to their root anatomy with diffuse-porous xylem structure in both beech and-opposing the stem structure-oak roots. In spruce, we found exclusively tracheids. We conclude that water can be redistributed within roots of different tree species along a moderate ψsoil gradient, accentuating HR as an important water source for drought-stressed plants, with potential implications for ecohydrological and plant physiological sciences. It remains to be shown to what extent HR occurs under field conditions in Central Europe.
Carbon (C) exuded via roots is proposed to increase under drought and facilitate important ecosystem functions. However, it is unknown how exudate quantities relate to the total C budget of a drought-stressed tree, that is, how much of net-C assimilation is allocated to exudation at the tree level.We calculated the proportion of daily C assimilation allocated to root exudation during early summer by collecting root exudates from mature Fagus sylvatica and Picea abies exposed to experimental drought, and combining above-and belowground C fluxes with leaf, stem and fine-root surface area.Exudation from individual roots increased exponentially with decreasing soil moisture, with the highest increase at the wilting point. Despite c. 50% reduced C assimilation under drought, exudation from fine-root systems was maintained and trees exuded 1.0% (F. sylvatica) to 2.5% (P. abies) of net C into the rhizosphere, increasing the proportion of C allocation to exudates two-to three-fold. Water-limited P. abies released two-thirds of its exudate C into the surface soil, whereas in droughted F. sylvatica it was only one-third.Across the entire root system, droughted trees maintained exudation similar to controls, suggesting drought-imposed belowground C investment, which could be beneficial for ecosystem resilience.
Hydraulic redistribution (HR) of soil water through plant roots is widely described; however its extent, especially in temperate trees, remains unclear. Here, we quantified HR of five temperate tree species. We hypothesized that both, HR within a plant and into the soil increase with higher water‐potential gradients, larger root conduit diameters and root‐xylem hydraulic conductivities as HR driving factors. Saplings of conifer (Picea abies, Pseudotsuga menziesii), diffuse‐porous (Acer pseudoplatanus) and ring‐porous species (Castanea sativa, Quercus robur) were planted in split‐root systems, where one plant had its roots split between two pots with different water‐potential gradients (0.23–4.20 MPa). We quantified HR via deuterium labelling. Species redistributed 0.39 ± 0.14 ml of water overnight (0.08 ± 0.01 ml/g root mass). Higher pre‐dawn water‐potential gradients, hydraulic conductivities and larger conduits significantly increased HR quantity. Hydraulic conductivity was the most important driving factor on HR amounts, within the plants (0.03 ± 0.01 ml/g) and into the soil (0.06 ± 0.01 ml/g). Additional factors as soil‐root contact should be considered, especially when calculating water transfer into the soil. Nevertheless, trees maintaining high‐xylem hydraulic conductivity showed higher HR amounts, potentially making them valuable ‘silvicultural tools’ to improve plant water status. A free Plain Language Summary can be found within the Supporting Information of this article.
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