Drying-rewetting cycles and γ-irradiation effects on enzyme activities of distinct layers from a Quercus ilex L. litter

In a Mediterranean climate, water stress is one of the principal constraints on proper forest ecosystem functioning. Drought influences rates of organic matter degradation by affecting microbial growth and enzyme activities. The objectives of this study were: (i) to evaluate the effect of repeated drying-rewetting cycles on cellulase, alkaline phosphatase and fluorescein diacetate (FDA) hydrolase activities of three distinct Quercus ilex L. litter layers, and (ii) to investigate the effect of these cycles on γ-irradiated litters in order to distinguish the abiotic influence on the fluctuations observed. Results, for all three layers, showed high correlations between litter water content and enzyme activities. Under mesocosm conditions, and using non-sterilized litter samples, cellulase, alkaline phosphatase, and FDA activities significantly decreased or increased during drying or rewetting cycles respectively. Significant differences were also found when evaluating the effect of litter depth on enzyme activities, the intermediate depth (OLv layer) generally being the most active. For γ-sterilized samples, FDA activity still fluctuated with drying-rewetting cycles. Assays showed that pre-humidification of γ-irradiated litter increased FDA activity two-fold in the first 30 min. All these results have shown that, following drying-rewetting cycles, some of the fluctuations occur independently of microbial growth, suggesting abiotic interactions, such as desorption, in combination with both solvatation status and conformational changes of enzymes.     

Does adding microbial mechanisms of decomposition improve soil organic matter models? A comparison of four models using data from a pulsed rewetting experiment

Contemporary soil organic matter (SOM) models have been successful at simulating decomposition across a range of spatial and temporal scales using first-order kinetics to represent the decomposition process; however, recent work suggests the simplicity of the first-order representation of decomposition is not adequate to capture the microbially-driven dynamics of SOM decomposition over short timescales. For example, the response of soils to drying-rewetting events may best be explained by microbial and/or exoenzyme controls on decomposition. To test if adding these microbial mechanisms improves the ability of SOM models to simulate the response of soils to short-term environmental changes, we developed four different SOM decomposition models with varying mechanistic complexity and compared their ability to simulate soil respiration from a pulsed drying-rewetting laboratory-based experiment. Specifically, we tested the ability of the models to capture the timing and magnitude of soil CO₂ efflux in response to rewetting or constant moisture conditions. The results of the comparison suggest that the inclusion of exoenzyme and microbial controls on decomposition can improve the ability to simulate pulsed rewetting dynamics; however, less mechanistic first-order models prevail under steady-state moisture conditions. These modeling results may have implications for understanding the long-term response of soil carbon stocks in response to local and regional climate change.     

干湿交替对土壤碳库和有机碳矿化的影响

水分是影响土壤活性碳库和惰性碳库周转过程的主导因子,而土壤有机碳的周转速率会对气候变化造成潜在的重要影响。以农田水稻土为供试土壤,通过培育试验研究了干湿交替过程对土壤有机碳矿化的影响,并利用两库叠加模型对土壤不同碳库及其降解动力学进行初步评估。结果表明:干湿交替激发了土壤呼吸,增加了土壤微生物代谢活性。三次湿润过程对土壤呼吸的激发量分别为119.3%、159.5%和87.3%,激发效应随干湿交替频率的增加先升高后降低。多次干湿交替后,土壤累积CO2释放量低于恒湿土壤,湿润所引起的激发的矿化量不足以弥补干旱期降低的矿化量。在湿润的数小时内,土壤溶解性有机碳含量先升高后降低。干湿交替提升了土壤活性碳库的降解速率,降低了惰性碳库的降解速率,湿润后土壤活性碳库显著增加。多次干湿交替降低了土壤真菌/细菌比,使土壤微生物群落结构发生变化,细菌成为优势种群。     

Soil microbial biomass and activity response to repeated drying-rewetting cycles along a soil fertility gradient modified by long-term fertilization management practices

The effects of repeated drying-rewetting (DRW) cycles on the microbial biomass and activity in soils taken from long-term field experiment plots with different fertilization (FERT) management practice histories were studied. We investigated the hypothesis that soil response to DRW cycles differs with soil fertility gradient modified by FERT management practices. The soils were incubated for 51 days, after exposure to either nine or three DRW cycles, or remaining at constant moisture content (CMC) at field capacity. We found that both DRW and FERT significantly affected soil properties including NH₄-N, NO₃-N, dissolved organic C (DOC), microbial biomass C (Cmic), basal soil respiration rate (BSR), urease activity (URE) and dehydrogenase activity (DHD). Except for NH₄-N and BSR, variation in the properties was largely explained by FERT, followed by DRW, and then their interaction. Irrespective of the soils' FERT treatment, repeated DRW cycles significantly raised the DOC and Cmic levels compared with CMC, and the DRW cycles also resulted in a significant decline in BSR and URE and increase in DHD, probably because the organisms were better-adapted to the drying and rewetting stresses. The variations in soil biological properties caused by DRW cycles showed a significantly negative relationship with the soil organic C content measured prior to the start of the DRW experiments, suggesting that soils with higher fertility are better able to maintain their original biological functions (i.e., have a higher functional stability) in response to DRW cycles.     

Soil CO2 efflux and extractable organic carbon fractions under simulated precipitation events in a Mediterranean Dehesa

The magnitude of CO₂ efflux pulses after rewetting a dry soil is highly variable and the factors regulating these pulses are poorly understood. In this field experiment, we aimed to study the C dynamics after simulated summer rainstorms in a Mediterranean open holm oak woodland (dehesa). We hypothesized that because the herbaceous cover is mostly dead during the summer in this ecosystem, the short-term CO₂ efflux (SR) after rewetting could mainly be explained by different measurable soil C fractions: i) K₂SO₄-extracted soil C (EOC); ii) microbial biomass C (MBC); or iii) chloroform-fumigated extracted C (CFE). On both grazed and abandoned dehesa sites, we simulated three summer rain events at two-week intervals and we measured SR discontinuously in three plots under tree canopy and in another three plots in open grassland. In each plot, C fractions and water content were estimated before (2 h) and after (36 h) each irrigation event. Following rewettings, SR increased up to ten times compared with non-irrigated plots. The CFE actually increased after rewetting in the first two irrigations but not in the third event, suggesting that the capacity of the soil to release labile organic C from soil aggregates or litter was reduced after each irrigation event. Overall, the C released as CO₂ in the first 24 h was related to the CFE existing before rewetting, which may help to explain the spatial variability in SR. However, the explained variability decreased after each irrigation, suggesting a change to a less labile composition of the CFE fraction as a consequence of multiple drying-rewetting cycles.     

Carbon pulses but not phosphorus pulses are related to decreases in microbial biomass during repeated drying and rewetting of soils

Drying and rewetting cycles are known to be important for the turnover of carbon (C) in soil, but less is known about the turnover of phosphorus (P) and its relation to C cycling. In this study the effects of repeated drying-rewetting (DRW) cycles on phosphorus (P) and carbon (C) pulses and microbial biomass were investigated. Soil (Chromic Luvisol) was amended with different C substrates (glucose, cellulose, starch; 2.5 g C kg⁻¹) to manipulate the size and community composition of the microbial biomass, thereby altering P mineralisation and immobilisation and the forms and availability of P. Subsequently, soils were either subjected to three DRW cycles (1 week dry/1 week moist) or incubated at constant water content (70% water filled pore space). Rewetting dry soil always produced an immediate pulse in respiration, between 2 and 10 times the basal rates of the moist incubated controls, but respiration pulses decreased with consecutive DRW cycles. DRW increased total CO₂ production in glucose and starch amended and non-amended soils, but decreased it in cellulose amended soil. Large differences between the soils persisted when respiration was expressed per unit of microbial biomass. In all soils, a large reduction in microbial biomass (C and P) occurred after the first DRW event, and microbial C and P remained lower than in the moist control. Pulses in extractable organic C (EOC) after rewetting were related to changes in microbial C only during the first DRW cycle; EOC concentrations were similar in all soils despite large differences in microbial C and respiration rates. Up to 7 mg kg⁻¹ of resin extractable P (P_resin) was released after rewetting, representing a 35-40% increase in P availability. However, the pulse in P_resin had disappeared after 7 d of moist incubation. Unlike respiration and reductions in microbial P due to DRW, pulses in P_resin increased during subsequent DRW cycles, indicating that the source of the P pulse was probably not the microbial biomass. Microbial community composition as indicated by fatty acid methyl ester (FAME) analysis showed that in amended soils, DRW resulted in a reduction in fungi and an increase in Gram-positive bacteria. In contrast, the microbial community in the non-amended soil was not altered by DRW. The non-selective reduction in the microbial community in the non-amended soil suggests that indigenous microbial communities may be more resilient to DRW. In conclusion, DRW cycles result in C and P pulses and alter the microbial community composition. Carbon pulses but not phosphorus pulses are related to changes in microbial biomass. The transient pulses in available P could be important for P availability in soils under Mediterranean climates.     

Characterization of the water soluble soil organic pool following the rewetting of dry soil in a drought-prone tallgrass prairie

To better understand the nature of the C flush that follows the rewetting of dry soil, we chemically characterized the water soluble pools following rewetting of soil dried to several different water potentials. To assess the impact that historical soil water status has on the size of the rewetting labile soluble pool, a laboratory water stress gradient was applied to soils that were collected from drought-prone and irrigated tallgrass prairie soils. In the laboratory, soils were either incubated at −33 kPa or dried steadily over a 0.6, 1, 2, or 3 day period to −1.5, −4, −15, and −45 MPa respectively. On the 4th day, samples were wetted back to −33 kPa and immediately assayed for soluble, microbial, or respiratory pools of carbon. After extraction, samples were also assayed using NMR, GC-MS, and LC-MS to assess carbohydrate, amino acid, osmolyte and sugar pools. The greater the degree of drying before rewetting was associated with greater concentrations of microbial, soluble and respiratory pools of carbon, increasing by 50, 400 and 250%, respectively, in the most water stressed compared to continuously moist soil. Compared to drought-prone soils, the amount of soluble C released as a result of rewetting was 30 to 50% greater in soils that were irrigated for 11 years. The pool of organics was not completely characterized and only small amounts of TBDMS and TMS derived compounds accounting for 2-4% of the soluble C pool were detected. In contrast, oligosaccharides constituted approximately 20-25% of the sample C. Our results suggest that the flush of C following wetting of a dry soil is not dominated by common microbial osmolytes (e.g. proline, glycine betaine, ectoine, glycerol, mannitol, trehalose). In light of this finding more research is needed to better understand the adaptations that microbial communities utilize to respond to the rewetting of dried soil.     

Response of microbial communities to water stress in irrigated and drought-prone tallgrass prairie soils

To better understand how water stress and availability affect the structure of microbial communities in soil, I measured the change in phospholipid fatty acids (PLFA) and the incorporation of ¹³C-labeled glucose into the PLFA following exposure to water stress. Overlaid on the laboratory water stress treatment, samples were collected from drought-prone and irrigated (11 years) tallgrass prairie soil (0-10 cm depth). In the laboratory, soils were either incubated at −250 kPa or dried steadily over a 3-d period to −45 MPa. On the fourth day, the dried samples were brought up to −250 kPa and then all samples received 250 μg of glucose-C (+4000 δ¹³C-PDB) solution that brought them to −33 kPa matric water potential. Samples were then extracted for PLFA following 6 and 24 h of incubation (25 °C). Non-metric multidimensional scaling (NMS) techniques and multi-response permutation procedure (MRPP) showed that the largest effect on the mol% distribution of PLFA was related to the field scale water addition experiment. In response to irrigation, the PLFA 16:1ω5, 18:1+, and 18:2ω6,9 showed increases, and a15:0, a17:0, and cy19:0 showed decreases in their respective mol%. Effects related to the induction of laboratory water stress were predominantly associated with a decrease in the mol% distribution of the putative fungal biomarker (18:2ω6,9) with little to no change in the mol% distribution of the bacterial biomarkers. Interestingly, the flow of C to the microbial community was not strongly related to any single PLFA, and differences were rather subtle, but multivariate MRPP detected change to the community structure related to the laboratory water stress treatment but not related to the 11 years of field irrigation. Our results suggest that both the total and the actively metabolizing bacterial community in soil were generally resistant to the effects of water stress brought by rewetting of dry soil. However, more research is needed to understand the nature of the fungal response to drying and rewetting in soil.