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Most empirical and modeling research on soil carbon (C) dynamics has focused on those processes that control and promote C stabilization. However, we lack a strong, generalizable understanding of the mechanisms through which soil organic carbon (SOC) is destabilized in soils. Yet a clear understanding of C destabilization processes in soil is needed to quantify the feedbacks of the soil C cycle to the Earth system. Destabilization includes processes that occur along a spectrum through which SOC shifts from a 'protected' state to an 'available' state to microbial cells where it can be mineralized to gaseous forms or to soluble forms that are then lost from the soil system. These processes fall into three general categories: (1) release from physical occlusion through processes such as tillage, bioturbation, or freeze-thaw and wetting-drying cycles; (2) C desorption from soil solids and colloids; and (3) increased C metabolism. Many processes that stabilize soil C can also destabilize C, and C gain or loss depends on the balance between competing reactions. For example, earthworms may both destabilize C through aggregate destruction, but may also create new aggregates and redistribute C into mineral horizon. Similarly, mycorrhizae and roots form new soil C but may also destabilize old soil C through priming and promoting microbial mining; labile C inputs cause C stabilization through increased carbon use efficiency or may fuel priming. Changes to the soil environment that affect the solubility of minerals or change the relative surfaces charges of minerals can destabilize SOC, including increased pH or in the reductive dissolution of Fe-bearing minerals. By considering these different physical, chemical, and biological controls as processes that contribute to soil C destabilization, we can develop thoughtful new hypotheses about the persistence and vulnerability of C in soils and make more accurate and robust predictions of soil C cycling in a changing environment.

Global forests are experiencing rising temperatures and more severe droughts, with consistently dire forecasts for negative future impacts. Current research on the physiological mechanisms underlying drought impacts is focused on the water- and carbon-associated mechanisms. The role of nutrients is notably missing from this research agenda. Here, we investigate what role, if any, forest nutrition plays for survival and recovery of forests during and after drought. High nutrient availability may play a detrimental role in drought survival due to preferential biomass allocation aboveground that (1) predispose plants to hydraulic constraints limiting photosynthesis and promoting hydraulic failure, (2) increases carbon costs during periods of carbon starvation, and (3) promote biotic attack due to low tissue carbon: nitrogen (C: N). When nutrient uptake occurs during drought, high nutrient availability can increase water use efficiency thus minimizing negative feedbacks between carbon and nutrient balance. Nutrients are released after drought ceases, which might promote faster recovery but the temporal dynamics of microbial immobilization and nutrient leaching have a significant impact on nutrient availability. We provide a framework for understanding nutrient impacts on drought survival that allows a more complete analysis of forest ecosystem responses.

The pulse of carbon dioxide (CO ₂) resulting from the first rainfall after the dry summer in Mediterranean ecosystems is so large that it is well documented at the landscape scale, with the CO ₂ released in a few days comparable in magnitude to the annual net carbon exchange of many terrestrial ecosystems. Although the origin of this CO ₂ is debated, we show that the pulse of CO ₂ is produced by a three-step resuscitation of the indigenous microbial community. Specific phylogenetic groups of microorganisms activate and contribute to the CO ₂ pulse at different times after a simulation of the first rainfall following the severe summer drought. Differential resuscitation was evident within 1 h of wet-up, with three primary response strategies apparent according to patterns of relative ribosomal quantity. Most bacteria could be classified as rapid responders (within 1 h of wet-up), intermediate responders (between 3 and 24 h after wet-up), or delayed responders (24–72 h after wet-up). Relative ribosomal quantities of rapid responders were as high in the prewet dry soils as at any other time, suggesting that specific groups of organisms may be poised to respond to the wet-up event, in that they preserve their capacity to synthesize proteins rapidly. Microbial response patterns displayed phylogenetic clustering and were primarily conserved at the subphylum level, suggesting that resuscitation strategies after wet-up of dry soil may be a phylogenetically conserved ecological trait.

Soil moisture controls microbial activity and soil carbon cycling. Because microbial activity decreases as soils dry, decomposition of soil organic matter (SOM) is thought to decrease with increasing drought length. Yet, microbial biomass and a pool of water-extractable organic carbon (WEOC) can increase as soils dry, perhaps implying microbes may continue to break down SOM even if drought stressed. Here, we test the hypothesis that WEOC increases as soils dry because exoenzymes continue to break down litter, while their products accumulate because they cannot diffuse to microbes. To test this hypothesis, we manipulated field plots by cutting off litter inputs and by irrigating and excluding precipitation inputs to extend or shorten the length of the dry season. We expected that the longer the soils would remain dry, the more WEOC would accumulate in the presence of litter, whereas shortening the length of the dry season, or cutting off litter inputs, would reduce WEOC accumulation. Lastly, we incubated grass roots in the laboratory and measured the concentration of reducing sugars and potential hydrolytic enzyme activities, strictly to understand the mechanisms whereby exoenzymes break down litter over the dry season. As expected, extending dry season length increased WEOC concentrations by 30% above the 108 μg C/g measured in untreated plots, whereas keeping soils moist prevented WEOC from accumulating. Contrary to our hypothesis, excluding plant litter inputs actually increased WEOC concentrations by 40% above the 105 μg C/g measured in plots with plants. Reducing sugars did not accumulate in dry senesced roots in our laboratory incubation. Potential rates of reducing sugar production by hydrolytic enzymes ranged from 0.7 to 10 μmol·g−1·h−1 and far exceeded the rates of reducing sugar accumulation (̃0.001 μmol·g−1·h−1). Our observations do not support the hypothesis that exoenzymes continue to break down litter to produce WEOC in dry soils. Instead, we develop the argument that physical processes are more likely to govern short-term WEOC dynamics via slaking of microaggregates that stabilize SOM and through WEOC redistribution when soils wet up, as well as through less understood effects of drought on the soil mineral matrix.

The rapid increase in microbial activity that occurs when a dry soil is rewetted has been well documented and is of great interest due to implications of changing precipitation patterns on soil C dynamics. Several studies have shown minor net changes in microbial population diversity or abundance following wet-up, but the gross population dynamics of bacteria and fungi resulting from soil wet-up are virtually unknown. Here we applied DNA stable isotope probing with H 2 18 O coupled with quantitative PCR to characterize new growth, survival, and mortality of bacteria and fungi following the rewetting of a seasonally dried California annual grassland soil. Microbial activity, as determined by CO 2 production, increased significantly within three hours of wet-up, yet new growth was not detected until after three hours, suggesting a pulse of nongrowth activity immediately following wet-up, likely due to osmo-regulation and resuscitation from dormancy in response to the rapid change in water potential. Total microbial abundance revealed little change throughout the seven-day post-wet incubation, but there was substantial turnover of both bacterial and fungal populations (49% and 52%, respectively). New growth was linear between 24 and 168 hours for both bacteria and fungi, with average growth rates of 2.3 × 10 8 bacterial 16S rRNA gene copies·[g dry mass] −1 ·h −1 and 4.3 × 10 7 fungal ITS copies·[g dry mass] −1 ·h −1 . While bacteria and fungi differed in their mortality and survival characteristics during the seven-day incubation, mortality that occurred within the first three hours was similar, with 25% and 27% of bacterial and fungal gene copies disappearing from the pre-wet community, respectively. The rapid disappearance of gene copies indicates that cell death, occurring either during the extreme dry down period (preceding five months) or during the rapid change in water potential due to wet-up, generates a significant pool of available C that likely contributes to the large pulse in CO 2 associated with wet-up. A dynamic assemblage of growing and dying organisms controlled the CO 2 pulse, but the balance between death and growth resulted in relatively stable total population abundances, even after a profound and sudden change in environment.

Wetting of dry soil triggers a pulse of microbial respiration that has been attributed to two broad mechanisms: (1) recycling of microbial cellular carbon (C), and (2) consumption of extracellular organic C made available to microbes by wetting. We evaluated these two mechanisms by measuring cumulative CO₂ release, changes in the size and chemical composition of microbial biomass, and water-extractable organic carbon (WEOC) concentrations following artificial wetting of soil sampled from two depths at each of seven sites across California spanning a range of geologic parent materials. In samples collected from surface soil (0–10 cm depth), we found that cumulative CO₂ release after wetting in the laboratory was most strongly correlated with microbial biomass. In these samples, the relative abundance of trehalose—a putative microbial osmolyte—decreased from 25% (SD = 12) to 16% (SD = 7) of the chloroform-labile fraction of the microbial biomass after wetting. This suggested a role for osmolyte consumption in generating the respiration pulse. In subsoil (40–50 cm depth, or sampled at contact with rock), however, the cumulative CO₂ release after wetting was unrelated to microbial biomass and more strongly related to WEOC. The concentrations of selected microbial biomass constituents (e.g. trehalose and amino acids) in WEOC were negligible (< 1%), suggesting that cell lysis was not important in generating WEOC in this study. The amount of WEOC relative to total organic C was greatest in subsoil, and negatively related to ammonium oxalate-extractable Fe (Pearson’s R = 0.42, p < 0.01), suggesting a role for soil mineralogical properties in controlling WEOC release. Together, these findings suggest that microbial cellular C and extracellular C jointly contribute to the respiration pulse, and that their relative contribution depends on depth.

Background and aim When soil is rewetted after drought, typically a transient pulse of mineralization and other microbial processes occur. This “Birch effect” translates into a temporarily elevated soil carbon dioxide efflux (SCE) and may alter nutrient availability. While rewetting effects on SCE have been frequently studied, effects on soil nutrient supply have rarely been considered despite potential relevance for plant nutrition during post-drought recovery. Methods We investigated the magnitude of the post-drought rewetting effect on SCE, ion exchange membrane-derived soil nutrient supply rates and leaf stoichiometry in a drought experiment in the Austrian Alps. We conducted the experiment on a managed grassland (MG) and a nearby abandoned grassland (AG). Results Under drought, soil moisture depleted faster at MG than at AG. Upon rewetting, the SCE pulse was significantly larger at MG than at AG, whereas N, P and K supplies were more strongly stimulated at AG. A large, transient rewetting effect on soil K supply (MG: +363 ± 132%; AG: +821 ± 195%) was reflected in elevated K in leaves of Leontodon hispidus . Conclusions Rewetting can alter post-drought nutrient availability in mountain grasslands, with particularly pronounced effects on soil K supply.

● The bacterial and fungal diversity decreased greater in 5%−36% DRW than 5%−25% DRW. ● Fungal network was complicated after 1-cycle DRW, but that for bacteria occurred until 4-cycle DRW. ● Stronger DRW treatment enhanced the pulse amplitude of respiration in soil. Altered drying-rewetting (DRW) procedures due to climate change may influence soil microbial properties and microbially-mediated carbon cycling in arid and semi-arid regions. However, the effects of DRW of different intensities on the microbial properties and respiration are not well understood. Thus, the responsive patterns of microbial communities and carbon mineralization in agriculture soil on the Chinese Loess Plateau to DRW treatments with different wetting intensities (5%−25% and 5%−36%) and frequency (1-cycle to 4-cycle) were investigated. Continuous moisture levels of 5%, 25% and 36% were used as control. Results revealed that the reduction of bacterial diversity and richness were greater for 5%−36% than 5%−25% treatment, while diversity of fungi was similar for different wetting intensities. Bacterial communities became clustered by wetting intensity rather than cycle number, however fungal community was unaffected by DRW. The complexity of bacterial co-occurrence network increased because of higher nodes, edges, average degree, diameter and average cluster coefficient after 4-cycles, and the interaction was more complex after 1-cycle for fungi. Rewetting caused a pulse-like increase of respiration rate, and the pulse amplitude was greater for DRW with high rewetting intensity and decreased with the increase of cycle number. The cumulative CO 2 emission for DRW treatments was lower than that for the continuous moisture conditions. The net reduction of carbon release for 5%−36% treatment was 1.18 times higher than that for 5%−25% treatment. Our study provides experimental evidence of the positive potential of DRW processes for maintaining soil carbon stock in an agriculture system on the Loess Plateau.

Tropical dry forests are already undergoing changes in the quantity and timing of rainfall, but there is great uncertainty over how these shifts will affect belowground carbon (C) cycling. While it has long been known that dry soils quickly release carbon dioxide (CO2) upon rewetting, the mechanisms underlying the so-called 'Birch effect' are still debated. Here, we quantified soil respiration pulses and their biotic predictors in response to simulated precipitation events in a regenerating tropical dry forest in Costa Rica. We also simulated the observed rewetting CO2 pulses with two soil carbon models: a conventional model assuming first-order decay rates of soil organic matter, and an enzyme-catalyzed model with Michaelis-Menten kinetics. We found that rewetting of dry soils produced an immediate and dramatic pulse of CO2, accompanied by rapid immobilization of nitrogen into the microbial biomass. However, the magnitude of the rewetting CO2 pulse was highly variable at fine spatial scales, and was well correlated with the size of the dissolved organic C pool prior to rewetting. Both the enzyme-catalyzed and conventional models were able to reproduce the Birch effect when respiration was coupled directly to microbial C uptake, although models differed in their ability to yield realistic estimates of SOC and microbial biomass pool sizes and dynamics. Our results suggest that changes in the timing and intensity of rainfall events in tropical dry forests will exert strong influence on ecosystem C balance by affecting the dynamics of microbial biomass growth.