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Traditionally, fine roots were grouped using arbitrary size categories, rarely capturing the heterogeneity in physiology, morphology and functionality among different fine root orders. Fine roots with different functional roles are rarely separated in microbiome‐focused studies and may result in confounding microbial signals and host‐filtering across different root microbiome compartments. Using a 26‐year‐old common garden, we sampled fine roots from four temperate tree species that varied in root morphology and sorted them into absorptive and transportive fine roots. The rhizoplane and rhizosphere were characterized using 16S rRNA gene and internal transcribed spacer region amplicon sequencing and shotgun metagenomics for the rhizoplane to identify potential microbial functions. Fine roots were subject to metabolomics to spatially characterize resource availability. Both fungi and bacteria differed according to root functional type. We observed additional differences between the bacterial rhizoplane and rhizosphere compartments for absorptive but not transportive fine roots. Rhizoplane bacteria, as well as the root metabolome and potential microbial functions, differed between absorptive and transportive fine roots, but not the rhizosphere bacteria. Functional differences were driven by sugar transport, peptidases and urea transport. Our data highlights the importance of root function when examining root‐microbial relationships, emphasizing different host selective pressures imparted on different root microbiome compartments. Summary Statement: Functionally discrete fine roots are often homogenized in microbiome studies. Here, we show that the microbial composition, potential microbial functions and the root metabolome differ with root functional type for four temperate tree species: Liriodendron tulipifera, Pinus strobus, Acer saccharum and Quercus rubra. [ABSTRACT FROM AUTHOR]

As climate change progresses, the frequency and duration of drought stress events are increasing. While the mechanisms of drought acclimation of trees has received considerable attention in recent years, the recovery processes remain critically understudied. We used a unique throughfall exclusion experiment in a mature temperate mixed forest consisting of the more isohydric Norway spruce and more anisohydric European beech, to study the recovery and resilience after drought release. We hypothesized that pre-dawn water potential (ΨPD) of both species will increase within 1 day after watering, while the recovery of stomatal conductance (g s) and the reversal of osmoregulation will be significantly delayed in the more isohydric spruce. Furthermore, we hypothesized that the xylem sap flow density (u daily) will not fully recover within the growing season due to the strong drought impact. After 5 years of summer drought, trees showed significantly reduced ΨPD, u daily and increased osmoregulation in leaves, but only isohydric spruce displayed increased leaf abscisic acid concentrations. In line with our hypothesis, ΨPD and g s recovered within 1 day in beech. Conversely, isohydric spruce showed delayed increases in ΨPD and g s. The delay in recovery of spruce was partially related to the replenishment of the stem water reservoir, as indicated by the missing response of u daily at the crown base compared with DBH level upon watering. However, u daily fully recovered only in the next growing season for beech and was still reduced in spruce. Nevertheless, in both species, osmotic acclimations of leaves were reversed within several weeks. While both species displayed full resilience to drought stress in water-related physiology, the recovery time was in several cases, e.g. u daily, ΨPD and g s, shorter for beech than for spruce. With future increases in the frequency of drought events under ongoing climate change, tree species that recover more quickly will be favored. [ABSTRACT FROM AUTHOR]

After drought events, tree recovery depends on sufficient carbon (C) allocation to the sink organs. The present study aimed to elucidate dynamics of tree‐level C sink activity and allocation of recent photoassimilates (Cnew) and stored C in c. 70‐year‐old Norway spruce (Picea abies) trees during a 4‐week period after drought release. We conducted a continuous, whole‐tree 13C labeling in parallel with controlled watering after 5 years of experimental summer drought. The fate of Cnew to growth and CO2 efflux was tracked along branches, stems, coarse‐ and fine roots, ectomycorrhizae and root exudates to soil CO2 efflux after drought release. Compared with control trees, drought recovering trees showed an overall 6% lower C sink activity and 19% less allocation of Cnew to aboveground sinks, indicating a low priority for aboveground sinks during recovery. In contrast, fine‐root growth in recovering trees was seven times greater than that of controls. However, only half of the C used for new fine‐root growth was comprised of Cnew while the other half was supplied by stored C. For drought recovery of mature spruce trees, in addition to Cnew, stored C appears to be critical for the regeneration of the fine‐root system and the associated water uptake capacity. [ABSTRACT FROM AUTHOR]

Summary: 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. [ABSTRACT FROM AUTHOR]

Hydraulic redistribution (HR) can buffer drought events of tree individuals, however, its relevance for neighbouring trees remains unclear. Here, we quantified HR to neighbouring trees in single‐ and mixed‐species combinations. We hypothesized that uptake of HR water positively correlates with root length, number of root tips and root xylem hydraulic conductivity and that neighbours in single‐species combinations receive more HR water than in phylogenetic distant mixed‐species combinations. In a split‐root experiment, a sapling with its roots split between two pots redistributed deuterium labelled water from a moist to a dry pot with an additional tree each. We quantified HR water received by the sapling in the dry pot for six temperate tree species. After 7 days, one quarter of the water in roots (2.1 ± 0.4 ml), stems (0.8 ± 0.2 ml) and transpiration (1.0 ± 0.3 ml) of the drought stressed sapling originated from HR. The amount of HR water transpired by the receiving plant stayed constant throughout the experiment. While the uptake of HR water increased with root length, species identity did not affect HR as saplings of Picea abies ((L.) Karst) and Fagus sylvatica (L.) in single‐ and mixed‐species combinations received the same amount of HR water. We quantified hydraulically redistributed water towards neighbouring saplings of six temperate tree species. One quarter of the water found in roots, shoots and transpiration originated from HR, with no differences due to species mixture, emphasizing the major importance of HR for temperate trees. [ABSTRACT FROM AUTHOR]

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 ± 26 m2 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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Hooves preserve the isotopic information laid down during their growth and may be used for reconstruction of animal feeding history. To assign certain positions along hooves to corresponding times, growth rates are required. Hoof growth rates are known for domestic animals; however, they cannot be obtained easily in wild animals. We estimated the hoof growth rate of the European roe deer (Capreolus capreolus L.) by using the immediate drop in δ13C along the hoof as a tag that is assigned to the date of maize (Zea mays L.) harvest. Keratin samples were taken each mm along 17 hooves and analyzed for their δ13C. A linear regression between (1) time differences of expected maize harvest to animal death and (2) distances between the points of the δ13C drop to the periople yielded the growth rate. Mean hoof growth rate was 0.122 mm/day (95% CI 0.014 mm/day) and 0.365%/day (±0.026%/day) of the hoof length and within the range of domestic animals. The method may be applied to determine growth rates of other incrementally growing tissues. Our estimated growth rate fosters dating isotopic information in hooves, facilitating research on feed resources and space use of roe deer. [ABSTRACT FROM AUTHOR]

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 ± 0.02MPa and -0.29 ± 0.03MPa), 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. [ABSTRACT FROM AUTHOR]