Forest understory plant and soil microbial response to an experimentally induced drought and heat‐pulse event: the importance of maintaining the continuum

Drought duration and intensity are expected to increase with global climate change. How changes in water availability and temperature affect the combined plant–soil–microorganism response remains uncertain. We excavated soil monoliths from a beech (Fagus sylvatica L.) forest, thus keeping the understory plant–microbe communities intact, imposed an extreme climate event, consisting of drought and/or a single heat‐pulse event, and followed microbial community dynamics over a time period of 28 days. During the treatment, we labeled the canopy with ¹³CO₂ with the goal of (i) determining the strength of plant–microbe carbon linkages under control, drought, heat and heat–drought treatments and (ii) characterizing microbial groups that are tightly linked to the plant–soil carbon continuum based on ¹³C‐labeled PLFAs. Additionally, we used 16S rRNA sequencing of bacteria from the Ah horizon to determine the short‐term changes in the active microbial community. The treatments did not sever within‐plant transport over the experiment, and carbon sinks belowground were still active. Based on the relative distribution of labeled carbon to roots and microbial PLFAs, we determined that soil microbes appear to have a stronger carbon sink strength during environmental stress. High‐throughput sequencing of the 16S rRNA revealed multiple trajectories in microbial community shifts within the different treatments. Heat in combination with drought had a clear negative effect on microbial diversity and resulted in a distinct shift in the microbial community structure that also corresponded to the lowest level of label found in the PLFAs. Hence, the strongest changes in microbial abundances occurred in the heat–drought treatment where plants were most severely affected. Our study suggests that many of the shifts in the microbial communities that we might expect from extreme environmental stress will result from the plant–soil–microbial dynamics rather than from direct effects of drought and heat on soil microbes alone.     

Coupled biogeochemical and hydrological responses of streams and rivers to drought

• 1 Severe or extreme droughts occurred about 10% of the time over a 105‐year record from central New Mexico, U.S.A., based on the Palmer Drought Severity Index.
• 2 Drought lowers water tables, creating extensive areas of groundwater recharge and fragmenting reaches of streams and rivers. Deeper groundwater inputs predominate as sources of surface flows during drought. Nutrient inputs to streams and rivers reflect the biogeochemistry of regional ground waters with longer subsurface residence times.
• 3 Inputs of bioavailable dissolved organic carbon to surface waters decrease during drought, with labile carbon limitation of microbial metabolism a byproduct of drought conditions.
• 4 Decreased inputs of organic forms of carbon, nitrogen and phosphorus and a decrease in the organic : inorganic ratio of nutrient inputs favours autotrophs over heterotrophs during drought.
• 5 The fate of autotrophic production during drought will be strongly influenced by the structure of the aquatic food web within impacted sites.

     

Key role of streambed moisture and flash storms for microbial resistance and resilience to long‐term drought

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Fast response of Scots pine to improved water availability reflected in tree‐ring width and δ13C

Drought‐induced forest decline, like the Scots pine mortality in inner‐Alpine valleys, will gain in importance as the frequency and severity of drought events are expected to increase. To understand how chronic drought affects tree growth and tree‐ring δ¹³C values, we studied mature Scots pine in an irrigation experiment in an inner‐Alpine valley. Tree growth and isotope analyses were carried out at the annual and seasonal scale. At the seasonal scale, maximum δ¹³C values were measured after the hottest and driest period of the year, and were associated with decreasing growth rates. Inter‐annual δ¹³C values in early‐ and latewood showed a strong correlation with annual climatic conditions and an immediate decrease as a response to irrigation. This indicates a tight coupling between wood formation and the freshly produced assimilates for trees exposed to chronic drought. This rapid appearance of the isotopic signal is a strong indication for an immediate and direct transfer of newly synthesized assimilates for biomass production. The fast appearance and the distinct isotopic signal suggest a low availability of old stored carbohydrates. If this was a sign for C‐storage depletion, an increasing mortality could be expected when stressors increase the need for carbohydrate for defence, repair or regeneration.     

Climate Change Affects Carbon Allocation to the Soil in Shrublands

Climate change may affect ecosystem functioning through increased temperatures or changes in precipitation patterns. Temperature and water availability are important drivers for ecosystem processes such as photosynthesis, carbon translocation, and organic matter decomposition. These climate changes may affect the supply of carbon and energy to the soil microbial population and subsequently alter decomposition and mineralization, important ecosystem processes in carbon and nutrient cycling. In this study, carried out within the cross-European research project CLIMOOR, the effect of climate change, resulting from imposed manipulations, on carbon dynamics in shrubland ecosystems was examined. We performed a ¹⁴C-labeling experiment to probe changes in net carbon uptake and allocation to the roots and soil compartments as affected by a higher temperature during the year and a drought period in the growing season. Differences in climate, soil, and plant characteristics resulted in a gradient in the severity of the drought effects on net carbon uptake by plants with the impact being most severe in Spain, followed by Denmark, with the UK showing few negative effects at significance levels of p 0.10. Drought clearly reduced carbon flow from the roots to the soil compartments. The fraction of the ¹⁴C fixed by the plants and allocated into the soluble carbon fraction in the soil and to soil microbial biomass in Denmark and the UK decreased by more than 60%. The effects of warming were not significant, but, as with the drought treatment, a negative effect on carbon allocation to soil microbial biomass was found. The changes in carbon allocation to soil microbial biomass at the northern sites in this study indicate that soil microbial biomass is a sensitive, early indicator of drought- or temperature-initiated changes in these shrubland ecosystems. The reduced supply of substrate to the soil and the response of the soil microbial biomass may help to explain the observed acclimation of CO₂ exchange in other ecosystems.     

Vegetation state changes in the course of shrub encroachment in an African savanna since about 1850 CE and their potential drivers

1. Shrub encroachment has far‐reaching ecological and economic consequences in many ecosystems worldwide. Yet, compositional changes associated with shrub encroachment are often overlooked despite having important effects on ecosystem functioning.
2. We document the compositional change and potential drivers for a northern Namibian Combretum woodland transitioning into a Terminalia shrubland. We use a multiproxy record (pollen, sedimentary ancient DNA, biomarkers, compound‐specific carbon (δ¹³C) and deuterium (δD) isotopes, bulk carbon isotopes (δ¹³Corg), grain size, geochemical properties) from Lake Otjikoto at high taxonomical and temporal resolution.
3. We provide evidence that state changes in semiarid environments may occur on a scale of one century and that transitions between stable states can span around 80 years and are characterized by a unique vegetation composition. We demonstrate that the current grass/woody ratio is exceptional for the last 170 years, as supported by n‐alkane distributions and the δ¹³C and δ¹³Corg records. Comparing vegetation records to environmental proxy data and census data, we infer a complex network of global and local drivers of vegetation change. While our δD record suggests physiological adaptations of woody species to higher atmospheric pCO₂ concentration and drought, our vegetation records reflect the impact of broad‐scale logging for the mining industry, and the macrocharcoal record suggests a decrease in fire activity associated with the intensification of farming. Impact of selective grazing is reflected by changes in abundance and taxonomical composition of grasses and by an increase of nonpalatable and trampling‐resistant taxa. In addition, grain‐size and spore records suggest changes in the erodibility of soils because of reduced grass cover.
4. Synthesis. We conclude that transitions to an encroached savanna state are supported by gradual environmental changes induced by management strategies, which affected the resilience of savanna ecosystems. In addition, feedback mechanisms that reflect the interplay between management legacies and climate change maintain the encroached state.

     

Defoliation alters the relative contributions of recent and non-recent assimilate to root exudation from Festuca rubra

The deposition of organic compounds from plant roots is a key determinant of rhizosphere microbial activity and community structure. Consequently, C-flow from roots to soil is an important process in coupling plant and microbial productivity, via impacts on microbial nutrient cycling in soil. Experimentally, isotopic tracers (¹³C or ¹⁴C) are used to track C inputs to soil and microbial communities. However, in many such studies the relationship between labelled C-flows and total C-flows are not established, limiting the interpretative value of the results. In this study, we applied steady-state near natural abundance ¹³CO₂ labelling to determine the impact of partial defoliation of Festuca rubra on root exudation. This approach in axenic culture facilitated determination of the contribution of pre- and post-defoliation assimilates both to root C-flow and plant tissues. The results demonstrated that total root exudation was increased in the two days following defoliation. This was concurrent with reduced net CO₂ assimilation and reduced allocation of post-defoliation assimilates below-ground and to active root meristems. Through determination of the δ¹³C of root exudates, it was established that the source of the increased root exudation was pre-defoliation assimilate. However, this response was transient, with reduced deposition of pre- and post-defoliation assimilates from roots during the period 2–4 d following defoliation. The results highlight the limitations of pulse-labelling approaches as a means of quantifying impacts of treatments on root exudation, particularly where the treatment is likely to affect plant C-partitioning or the balance between deposition to, and re-mobilization from, C-storage pools.     

Use of decreasing foliar carbon isotope discrimination during water limitation as a carbon tracer to study whole plant carbon allocation

Foliar carbon isotope discrimination (Δ) of C₃ plants decreases in water‐deficit situations as discrimination by the photosynthetic primary carboxylation reaction decreases. This diminished Δ in leaves under water deficit can be used as a tracer to study whole plant carbon allocation patterns. Carbon isotope composition (δ¹³C value) of leaf hot water extracts or leaf tissue sap represents a short‐term integral of leaf carbon isotope discrimination and thus represents the δ¹³C value of source carbon that may be distributed within a plant in water‐deficit situations. By plotting the δ¹³C values of source carbon against the δ¹³C values of sink tissues, such as roots or stems, it is possible to assess carbon allocation to and incorporation into sink organs in relation to already present biomass. This natural abundance labelling method has been tested in three independent experiments, a one‐year field study with the fruit tree species Ziziphus mauritiana and peach (Prunus persica), a medium‐term drought stress experiment with Ziziphus rotundifolia trees in the glasshouse, and a short‐term drought stress experiment with soybean (Glycine max). The data show that the natural abundance labelling method can be applied to qualitatively assess carbon allocation in drought‐stressed plants. Although it is not possible to estimate exact fluxes of assimilated carbon during water deficit the method represents an easy to use tool to study integrated plant adaptations to drought stress. In addition, it is a less laborious method that can be applied in field studies as well as in controlled experiments, with plants from any developmental stage.     

Environmental controls on benthic food web functions and carbon resource use in subarctic lakes

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Partitioning pattern of carbon flux in a Kobresia grassland on the Qinghai‐Tibetan Plateau revealed by field 13C pulse‐labeling

Characterizing the carbon turnover in terrestrial ecosystems is critical for understanding and predicting carbon dynamics in ecosystems. We used in situ¹³C pulse labeling to track photosynthetic carbon fluxes from shoot to roots and to soil in a Kobresia humilis meadow on the Qinghai‐Tibet Plateau. We found that about 36.7% of labeled carbon was translocated out from the shoots within the first 24 h after photosynthetic uptake. This is equivalent to 66.1% of total ¹³C moving out from the shoot during the 32‐day chase period, indicating a rapid and large translocation of newly fixed carbon to belowground parts in these alpine plants. 58.7% of the assimilated ¹³C was transferred belowground. At the end of the chase phase, 30.9% was retained in living roots, 3.4% in dead roots, 17.2% lost as belowground respiration and 7.3% remained in the soil. In the four carbon pools (i.e., shoots, living roots, dead roots, and soil pools), living roots consistently had the highest proportion of ¹³C in the plant–soil system during the 32 days. Based on the ¹³C partitioning pattern and biomass production, we estimate a total of 4930 kg C ha^?1 was allocated belowground during the vegetation growth season in this alpine meadow. Of this, roots accumulated 2868 kg C ha^?1 and soils accumulated 613 kg C ha^?1. This study suggests that carbon storage in belowground carbon pools plays the most important role in carbon cycles in the alpine meadow.     

The effects of drought and herbivory on plant–herbivore interactions across 16 soybean genotypes in a field experiment

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Drought decreases incorporation of recent plant photosynthate into soil food webs regardless of their trophic complexity

Theory suggests that more complex food webs promote stability and can buffer the effects of perturbations, such as drought, on soil organisms and ecosystem functions. Here, we tested experimentally how soil food web trophic complexity modulates the response to drought of soil functions related to carbon cycling and the capture and transfer below‐ground of recent photosynthate by plants. We constructed experimental systems comprising soil communities with one, two or three trophic levels (microorganisms, detritivores and predators) and subjected them to drought. We investigated how food web trophic complexity in interaction with drought influenced litter decomposition, soil CO₂ efflux, mycorrhizal colonization, fungal production, microbial communities and soil fauna biomass. Plants were pulse‐labelled after the drought with ¹³C‐CO₂ to quantify the capture of recent photosynthate and its transfer below‐ground. Overall, our results show that drought and soil food web trophic complexity do not interact to affect soil functions and microbial community composition, but act independently, with an overall stronger effect of drought. After drought, the net uptake of ¹³C by plants was reduced and its retention in plant biomass was greater, leading to a strong decrease in carbon transfer below‐ground. Although food web trophic complexity influenced the biomass of Collembola and fungal hyphal length, ¹³C enrichment and the net transfer of carbon from plant shoots to microbes and soil CO₂ efflux were not affected significantly by varying the number of trophic groups. Our results indicate that drought has a strong effect on above‐ground–below‐ground linkages by reducing the flow of recent photosynthate. Our results emphasize the sensitivity of the critical pathway of recent photosynthate transfer from plants to soil organisms to a drought perturbation, and show that these effects may not be mitigated by the trophic complexity of soil communities, at least at the level manipulated in this experiment.     

Beaver activity increases aquatic subsidies to terrestrial consumers

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Impact of interspecific competition and drought on the allocation of new assimilates in trees

In trees, the interplay between reduced carbon assimilation and the inability to transport carbohydrates to the sites of demand under drought might be one of the mechanisms leading to carbon starvation. However, we largely lack knowledge on how drought effects on new assimilate allocation differ between species with different drought sensitivities and how these effects are modified by interspecific competition. We assessed the fate of ¹³C labelled assimilates in above‐ and belowground plant organs and in root/rhizosphere respired CO₂ in saplings of drought‐tolerant Norway maple (Acer platanoides) and drought‐sensitive European beech (Fagus sylvatica) exposed to moderate drought, either in mono‐ or mixed culture. While drought reduced stomatal conductance and photosynthesis rates in both species, both maintained assimilate transport belowground. Beech even allocated more new assimilate to the roots under moderate drought compared to non‐limited water supply conditions, and this pattern was even more pronounced under interspecific competition. Even though maple was a superior competitor compared to beech under non‐limited soil water conditions, as indicated by the changes in above‐ and belowground biomass of both species in the interspecific competition treatments, we can state that beech was still able to efficiently allocate new assimilate belowground under combined drought and interspecific competition. This might be seen as a strategy to maintain root osmotic potential and to prioritise root functioning. Our results thus show that beech tolerates moderate drought stress plus competition without losing its ability to supply belowground tissues. It remains to be explored in future work if this strategy is also valid during long‐term drought exposure.     

Aphids in a changing world: testing the plant stress,plant vigour and pulsed stress hypotheses

• 1 The plant stress, plant vigour and pulsed stress hypotheses describe the relationships between drought stress, plant quality and herbivore performance. We used an aphid‐Brassica system to test these hypotheses under different drought treatments.
• 2 The quantity of water added per plant/week was 75%, 50% and 25% of the control (unstressed) water regime for low, medium and high drought stress, respectively, and 50% applied fortnightly for pulsed drought stress. The performance of a ‘senescence’ (generalist) and a ‘flush’ feeder (specialist) aphid species and host plant quality were assessed.
• 3 Drought treatments had a similar effect on the fecundity and intrinsic rate of increase of both aphid species. Aphid performance on unstressed and highly drought‐stressed plants was significantly lower compared with medium drought stress. On average, 20% greater fecundity and 40% greater intrinsic rates of increase were recorded for both aphid species at medium drought stress compared with unstressed plants.
• 4 Plant biomass and relative water contents were significantly greater for unstressed plants compared with high and pulsed drought treatments. Foliar nitrogen concentration was significantly greater in the high drought stress and pulsed treatments, and the dominant glucosinolate (glucobrassicin) concentration was significantly greater in drought stress treatments.
• 5 The present study supports the plant stress hypothesis, although the plant vigour and pulsed stress hypotheses are not supported by our data. The implications of these findings for plant–herbivore interactions under changing environmental conditions are discussed.

     

Effects of simulated drought on the carbon balance of Everglades short‐hydroperiod marsh

Hydrology drives the carbon balance of wetlands by controlling the uptake and release of CO₂ and CH₄. Longer dry periods in between heavier precipitation events predicted for the Everglades region, may alter the stability of large carbon pools in this wetland's ecosystems. To determine the effects of drought on CO₂ fluxes and CH₄ emissions, we simulated changes in hydroperiod with three scenarios that differed in the onset rate of drought (gradual, intermediate, and rapid transition into drought) on 18 freshwater wetland monoliths collected from an Everglades short‐hydroperiod marsh. Simulated drought, regardless of the onset rate, resulted in higher net CO₂ losses net ecosystem exchange (NEE) over the 22‐week manipulation. Drought caused extensive vegetation dieback, increased ecosystem respiration (R_eco), and reduced carbon uptake gross ecosystem exchange (GEE). Photosynthetic potential measured by reflective indices (photochemical reflectance index, water index, normalized phaeophytinization index, and the normalized difference vegetation index) indicated that water stress limited GEE and inhibited R_eco. As a result of drought‐induced dieback, NEE did not offset methane production during periods of inundation. The average ratio of net CH₄ to NEE over the study period was 0.06, surpassing the 100‐year greenhouse warming compensation point for CH₄ (0.04). Drought‐induced diebacks of sawgrass (C₃) led to the establishment of the invasive species torpedograss (C₄) when water was resupplied. These changes in the structure and function indicate that freshwater marsh ecosystems can become a net source of CO₂ and CH₄ to the atmosphere, even following an extended drought. Future changes in precipitation patterns and drought occurrence/duration can change the carbon storage capacity of freshwater marshes from sinks to sources of carbon to the atmosphere. Therefore, climate change will impact the carbon storage capacity of freshwater marshes by influencing water availability and the potential for positive feedbacks on radiative forcing.     

Water‐level fluctuations affect sediment properties,carbon flux and growth of the isoetid Littorella uniflora in oligotrophic lakes

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Fire deficits have increased drought sensitivity in dry conifer forests: Fire frequency and tree‐ring carbon isotope evidence from Central Oregon

A century of fire suppression across the Western United States has led to more crowded forests and increased competition for resources. Studies of forest thinning or stand conditions after mortality events have provided indirect evidence for how competition can promote drought stress and predispose forests to severe fire and/or bark beetle outbreaks. Here, we demonstrate linkages between fire deficits and increasing drought stress through analyses of annually resolved tree‐ring growth, fire scars, and carbon isotope discrimination (Δ¹³C) across a dry mixed‐conifer forest landscape. Fire deficits across the study area have increased the sensitivity of leaf gas exchange to drought stress over the past >100 years. Since 1910, stand basal area in these forests has more than doubled and fire‐return intervals have increased from 25 to 140 years. Meanwhile, the portion of interannual variation in tree‐ring Δ¹³C explained by the Palmer Drought Severity Index has more than doubled in ca. 300–500‐year‐old Pinus ponderosa as well as in fire‐intolerant, ca. 90–190‐year‐old Abies grandis. Drought stress has increased in stands with a basal area of ≥25 m²/ha in 1910, as indicated by negative temporal Δ¹³C trends, whereas stands with basal area ≤25 m²/ha in 1910, due to frequent or intense wildfire activity in decades beforehand, were initially buffered from increased drought stress and have benefited more from rising ambient carbon dioxide concentrations, [CO₂], as demonstrated by positive temporal Δ¹³C trends. Furthermore, the average Δ¹³C response across all P. ponderosa since 1830 indicates that photosynthetic assimilation rates and stomatal conductance have been reduced by ~10% and ~20%, respectively, compared to expected trends due to increasing [CO₂]. Although disturbance legacies contribute to local‐scale intensity of drought stress, fire deficits have reduced drought resistance of mixed‐conifer forests and made them more susceptible to challenges by pests and pathogens and other disturbances.     

Tracking a century of change in trophic structure and dynamics in a floodplain wetland: integrating palaeoecological and palaeoisotopic evidence

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