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Drought is a severe natural risk that increases drying-rewetting frequencies of soils. Yet, it remains largely unknown how forest ecosystems respond to dry-wet cycles, hampering our ability to evaluate the overall sink and source functionality for this large carbon pool. Recent investigations suggest that the release of soluble carbon via root exudation increases under drought, influencing soil carbon stabilization and mineralization. However, an integration of root exudation into the carbon allocation dynamics of drought stressed trees is missing. We hypothesized that roots in dry soil layers have a higher exudation rate than roots in more moist layers across different soil depths. Further, we tested if higher exudation rates under drought are attenuated by reduced root abundance in dry soils and if the fraction of root exudation from total carbon allocation increases with decreasing photosynthesis rates under drought. At the KROOF experimental site in southern Germany, where mature beech (Fagus sylvatica L.) and spruce (Picea abies (L.) Karst.) trees were exposed to artificial drought stress for five consecutive growing seasons, we show that at the root level root exudation rate increases in drier soils. Especially roots in the upper soil profile and roots of spruce trees increased root exudation under drought. When scaled to whole tree level, we did not find differences in total exudation between drought stressed and control trees, indicating sustained root exudation at the tree level under drought. As photosynthesis rates and therefore total carbon assimilation was substantially reduced under drought (by 50 % in beech and almost 70 % in spruce), the fraction of root exudation from total assimilation slightly increased for drought stressed trees. Our results demonstrate that stimulation of root exudation rates with drought exists in natural temperate forest ecosystems but might be mitigated by reduced fine root abundance under drought. Nevertheless, increased exudation per root surface area will have localized impacts on rhizosphere microbial composition and activity especially in the topsoil exposed to more extreme dry-wet cycles. Finally, also the exudate composition can help to determine how priming of soil organic matter relates to belowground carbon allocation dynamics and to disclose processes of complementary species interaction and should be emphasised in future studies.

This contribution presents the result of a free-air 13C labeling experiment on mature Norway spruce (P. abies [L.] KARST.) upon watering after five years of recurrent summer drought in southern Germany, focusing on whole tree allocation processes. Mature spruce trees had been exposed to recurrent summer drought from 2014 to 2018 through complete exclusion of precipitation throughfall from spring to late fall (i.e., March to November). In early summer 2019, the drought stressed spruce trees were watered to investigate their recovery processes. In parallel with the watering, we conducted a whole-tree 13C labeling in canopies and traced the signal in various C sinks, i.e. stem phloem and CO2 efflux, tree rings at different heights, coarse roots, fine root tips, mycorrhiza, root exudates, and soil CO2 efflux.We hypothesize that drought stressed spruce preferentially allocates newly assimilated C to belowground sinks upon drought release. Conversely to our expectations, allocation to belowground C sinks was not stimulated in drought stressed compared to control spruce. Likewise, the relative amount of recently fixed C allocated to aboveground sinks did not differ between treatments. Our findings suggest that the belowground C sinks are not of higher priority for the allocation of newly assimilated C upon watering after long-term drought. The observed allocation pattern is discussed taking total above- and belowground biomass as well as C source/sink relations into account.

Ground-level ozone (O3) is a widespread air pollutant causing extensive injuries in plants. However, its effects on perennial energy crops remain poorly understood due to technical difficulties in cultivating fast-growing shrubs for biomass production under O3 treatment on the field. Here we present the results of a two-year evaluation in the framework of which willow (Salix sachalinensis F. Schmid) shrubs were exposed to ambient (AOZ) or elevated (EOZ) O3 in two successive growing seasons (2014, 2015) and treated with 0 (EDU0) or 400 mg L−1 (EDU400) ethylenediurea spray in the second growing season. In 2014, EOZ altered the chemical composition of both top young and fallen leaves, and a novel mechanism of decreasing Mg in fallen leaves while highly enriching it in young top leaves was revealed in shrubs exposed to EOZ. In 2015, EDU400 alleviated EOZ-induced decreases in leaf fresh mass to dry mass ratio (FM/DM) and leaf mass per area (LMA). While EDU400 protected against EOZ-induced suppression of the maximum rate at which leaves can fix carbon (Amax) in O3-asymptomatic leaves, it did not alleviate EOZ-induced suppression of the maximum rates of carboxylation (VCmax) and electron transport (Jmax) and chlorophylls a, b, and a + b in the same type of leaves. In O3-symptomatic leaves, however, EDU400 alleviated EOZ-induced suppression of chlorophylls a and a + b, indicating different mode of action of EDU between O3-asymptomatic and O3-symptomatic leaves. Extensive herbivory occurred only in AOZ-exposed plants, leading to suppressed biomass production, while EOZ also led to a similar suppression of biomass production (EDU0 × EOZ vs. EDU400 × EOZ). In 2016, carry-over effects were also evaluated following cropping and transplantation into new ambient plots. Effects of EOZ in the preceding growing seasons extended to the third growing season in the form of suppressed ratoon biomass production, indicating carry-over effect of EOZ. Although EDU400 protected against EOZ-induced suppression of biomass production when applied in 2015, there was no carry-over effect of EDU in the absence of EDU treatment in 2016. The results of this study provide novel mechanistic understandings of O3 and EDU modes of action and can enlighten cultivation of willow as energy crop.