The effects of tree rhizodeposition on soil exoenzyme activity, dissolved organic carbon, and nutrient availability in a subalpine forest ecosystem

Previous studies have found that root carbon inputs to the soil can stimulate the mineralization of existing soil carbon (C) pools. It is still uncertain, however, whether this “primed” C is derived from elevated rates of soil organic matter (SOM) decomposition, greater C release from microbial pools, or both. The goal of this research was to determine how the activities of the microbial exoenzymes that control SOM decomposition are affected by root C inputs. This was done by manipulating rhizodeposition with tree girdling in a coniferous subalpine forest in the Rocky Mountains of Colorado, USA, and following changes in the activities of nine exoenzymes involved in decomposition, as well as soil dissolved organic C, dissolved organic and inorganic nitrogen (N), and microbial biomass C and N. We found that rhizodeposition is high in the spring, when the soils are still snow-covered, and that there are large ephemeral populations of microorganisms dependent upon this C. Microbial N acquisition from peptide degradation increased with increases in microbial biomass when rhizodeposition was highest. However, our data indicate that the breakdown of cellulose, lignin, chitin, and organic phosphorus are not affected by springtime increases in soil microbial biomass associated with increases in rhizodeposition. We conclude that the priming of soil C mineralization by rhizodeposition is due to growth of the microbial biomass and an increase in the breakdown of N-rich proteins, but not due to increases in the degradation of plant litter constituents such as cellulose and lignin.     

Quantitative assessment of microbial necromass contribution to soil organic matter

Soil carbon transformation and sequestration have received significant interest in recent years due to a growing need for quantitating its role in mitigating climate change. Even though our understanding of the nature of soil organic matter has recently been substantially revised, fundamental uncertainty remains about the quantitative importance of microbial necromass as part of persistent organic matter. Addressing this uncertainty has been hampered by the absence of quantitative assessments whether microbial matter makes up the majority of the persistent carbon in soil. Direct quantitation of microbial necromass in soil is very challenging because of an overlapping molecular signature with nonmicrobial organic carbon. Here, we use a comprehensive analysis of existing biomarker amino sugar data published between 1996 and 2018, combined with novel appropriation using an ecological systems approach, elemental carbon–nitrogen stoichiometry, and biomarker scaling, to demonstrate a suit of strategies for quantitating the contribution of microbe‐derived carbon to the topsoil organic carbon reservoir in global temperate agricultural, grassland, and forest ecosystems. We show that microbial necromass can make up more than half of soil organic carbon. Hence, we suggest that next‐generation field management requires promoting microbial biomass formation and necromass preservation to maintain healthy soils, ecosystems, and climate. Our analyses have important implications for improving current climate and carbon models, and helping develop management practices and policies.     

Soil Carbon, Nitrogen and Phosphorus Dynamics as Affected by Solarization Alone or Combined with Organic Amendment

Soil solarization, alone or combined with organic amendment, is an increasingly attractive approach for managing soil-borne plant pathogens in agricultural soils. Even though it consists in a relatively mild heating treatment, the increased soil temperature may strongly affect soil microbial processes and nutrients dynamics. This study aimed to investigate the impact of solarization, either with or without addition of farmyard manure, in soil dynamics of various C, N and P pools. Changes in total C, N and P contents and in some functionally-related labile pools (soil microbial biomass C and N, K₂SO₄-extractable C and N, basal respiration, KCl-exchangeable ammonium and nitrate, and water-soluble P) were followed across a 72-day field soil solarization experiment carried out during a summer period on a clay loam soil in Southern Italy. Soil physico-chemical properties (temperature, moisture content and pH) were also monitored. The average soil temperature at 8-cm depth in solarized soils approached 55 °C as compared to 35 °C found in nonsolarized soil. Two-way ANOVA (solarization×organic amendment) showed that both factors significantly affected most of the above variables, being the highest influence exerted by the organic amendment. With no manure addition, solarization did not significantly affect soil total C, N and P pools. Whereas soil pH, microbial biomass and, at a greater extent, K₂SO₄-extractable N and KCl-exchangeable ammonium were greatly affected. An increased release of water-soluble P was also found in solarized soils. Yet, solarization altered the quality of soluble organic residues released in soil as it lowered the C-to-N ratio of both soil microbial biomass and K₂SO₄-extractable organic substrates. Additionally, in solarized soils the metabolic quotient (qCO₂) significantly increased while the microbial biomass C-to-total organic C ratio (microbial quotient) decreased over the whole time course. We argued that soil solarization promoted the mineralization of readily decomposable pools of the native soil organic matter (e.g. the microbial biomass) thus rendering larger, at least over a short-term, the available fraction of some soil mineral nutrients, namely N and P forms. However, over a longer prospective solarization may lead to an over-exploitation of labile organic resources in agricultural soils. Manure addition greatly increased the levels of both total and labile C, N and P pools. Thus, addition of organic amendments could represent an important strategy to protect agricultural lands from excessive soil resources exploitation and to maintain soil fertility while enhancing pest control.     

Microbial necromass in cropland soils: A global meta-analysis of management effects

Microbial necromass is a large and persistent component of soil organic carbon (SOC), especially under croplands. The effects of cropland management on microbial necromass accumulation and its contribution to SOC have been measured in individual studies but have not yet been summarized on the global scale. We conducted a meta-analysis of 481-paired measurements from cropland soils to examine the management effects on microbial necromass and identify the optimal conditions for its accumulation. Nitrogen fertilization increased total microbial necromass C by 12%, cover crops by 14%, no or reduced tillage (NT/RT) by 20%, manure by 21%, and straw amendment by 21%. Microbial necromass accumulation was independent of biochar addition. NT/RT and straw amendment increased fungal necromass and its contribution to SOC more than bacterial necromass. Manure increased bacterial necromass higher than fungal, leading to decreased ratio of fungal-to-bacterial necromass. Greater microbial necromass increases after straw amendments were common under semi-arid and in cool climates in soils with pH <8, and were proportional to the amount of straw input. In contrast, NT/RT increased microbial necromass mainly under warm and humid climates. Manure application increased microbial necromass irrespective of soil properties and climate. Management effects were especially strong when applied during medium (3–10 years) to long (10+ years) periods to soils with larger initial SOC contents, but were absent in sandy soils. Close positive links between microbial biomass, necromass and SOC indicate the important role of stabilized microbial products for C accrual. Microbial necromass contribution to SOC increment (accumulation efficiency) under NT/RT, cover crops, manure and straw amendment ranged from 45% to 52%, which was 9%–16% larger than under N fertilization. In summary, long-term cropland management increases SOC by enhancing microbial necromass accumulation, and optimizing microbial necromass accumulation and its contribution to SOC sequestration requires site-specific management.     

Incorporation of Plant Residues into Soil Organic Matter Fractions With Grassland Management Practices in the North American Midwest

Disturbed grassland soils are often cited as having the potential to store large amounts of carbon (C). Fertilization of grasslands can promote soil C storage, but little is known about the generation of recalcitrant pools of soil organic matter (SOM) with management treatments, which is critical for long-term soil C storage. We used a combination of soil incubations, size fractionation and acid hydrolysis of SOM, [C], [N], and stable isotopic analyses, and biomass quality indices to examine how fertilization and haying can impact SOM dynamics in Kansan grassland soils. Fertilized soils possessed 113% of the C possessed by soils subjected to other treatments, an increase predominantly harbored in the largest size fraction (212–2,000 μm). This fraction is frequently associated with more labile material. Haying and fertilization/haying, treatments that more accurately mimic true management techniques, did not induce any increase in soil C. The difference in ¹⁵N-enrichment between size fractions was consistent with a decoupling of SOM processing between pools with fertilization, congruent with gains of SOM in the largest size fraction promoted by fertilization not moving readily into smaller fractions that frequently harbor more recalcitrant material. Litterfall and root biomass C inputs increased 104% with fertilization over control plots, and this material possessed lower C:N ratios. Models of incubation mineralization kinetics indicate that fertilized soils have larger pools of labile organic C. Model estimates of turnover rates of the labile and recalcitrant C pools did not differ between treatments (65.5 ± 7.2 and 2.9 ± 0.3 μg C d⁻¹, respectively). Although fertilization may promote greater organic inputs into these soils, much of that material is transformed into relatively labile forms of soil C; these data highlight the challenges of managing grasslands for long-term soil C sequestration.     

Microbial necromass under global change and implications for soil organic matter

Microbial necromass is an important source and component of soil organic matter (SOM), especially within the most stable pools. Global change factors such as anthropogenic nitrogen (N), phosphorus (P), and potassium (K) inputs, climate warming, elevated atmospheric carbon dioxide (eCO₂), and periodic precipitation reduction (drought) strongly affect soil microorganisms and consequently, influence microbial necromass formation. The impacts of these global change factors on microbial necromass are poorly understood despite their critical role in the cycling and sequestration of soil carbon (C) and nutrients. Here, we conducted a meta-analysis to reveal general patterns of the effects of nutrient addition, warming, eCO₂, and drought on amino sugars (biomarkers of microbial necromass) in soils under croplands, forests, and grasslands. Nitrogen addition combined with P and K increased the content of fungal (+21%), bacterial (+22%), and total amino sugars (+9%), consequently leading to increased SOM formation. Nitrogen addition alone increased solely bacterial necromass (+10%) because the decrease of N limitation stimulated bacterial more than fungal growth. Warming increased bacterial necromass, because bacteria have competitive advantages at high temperatures compared to fungi. Other global change factors (P and NP addition, eCO₂, and drought) had minor effects on microbial necromass because of: (i) compensation of the impacts by opposite processes, and (ii) the short duration of experiments compared to the slow microbial necromass turnover. Future studies should focus on: (i) the stronger response of bacterial necromass to N addition and warming compared to that of fungi, and (ii) the increased microbial necromass contribution to SOM accumulation and stability under NPK fertilization, and thereby for negative feedback to climate warming.     

Experimental litterfall manipulation drives large and rapid changes in soil carbon cycling in a wet tropical forest

Global changes such as variations in plant net primary production are likely to drive shifts in leaf litterfall inputs to forest soils, but the effects of such changes on soil carbon (C) cycling and storage remain largely unknown, especially in C‐rich tropical forest ecosystems. We initiated a leaf litterfall manipulation experiment in a tropical rain forest in Costa Rica to test the sensitivity of surface soil C pools and fluxes to different litter inputs. After only 2 years of treatment, doubling litterfall inputs increased surface soil C concentrations by 31%, removing litter from the forest floor drove a 26% reduction over the same time period, and these changes in soil C concentrations were associated with variations in dissolved organic matter fluxes, fine root biomass, microbial biomass, soil moisture, and nutrient fluxes. However, the litter manipulations had only small effects on soil organic C (SOC) chemistry, suggesting that changes in C cycling, nutrient cycling, and microbial processes in response to litter manipulation reflect shifts in the quantity rather than quality of SOC. The manipulation also affected soil CO ₂ fluxes; the relative decline in CO ₂ production was greater in the litter removal plots (?22%) than the increase in the litter addition plots (+15%). Our analysis showed that variations in CO ₂ fluxes were strongly correlated with microbial biomass pools, soil C and nitrogen (N) pools, soil inorganic P fluxes, dissolved organic C fluxes, and fine root biomass. Together, our data suggest that shifts in leaf litter inputs in response to localized human disturbances and global environmental change could have rapid and important consequences for belowground C storage and fluxes in tropical rain forests, and highlight differences between tropical and temperate ecosystems, where belowground C cycling responses to changes in litterfall are generally slower and more subtle.     

Impacts of chronic nitrogen additions vary seasonally and by microbial functional group in tundra soils

Previous studies have shown that fertilization with nitrogen depresses overall microbial biomass and activity in soil. In the present study we broaden our understanding of this phenomenon by studying the seasonality of responses of specific microbial functional groups to chronic nitrogen additions in alpine tundra soils. We measured soil enzyme activities, mineralization kinetics for 8 substrates, biomass of 8 microbial functional groups, and changes in N and carbon pools in the soil. Our approach allowed us to compare the ability of the soil microbial biomass to utilize various substrates in addition to allowing us to estimate changes in biomass of microbial functional groups that are involved in carbon and nitrogen cycling. Overall microbial activity and biomass was reduced in fertilized plots, whereas pools of N in the soil and microbial biomass N were higher in fertilized plots. The negative effects of N were most prominent in the summer. Biomass of the dominant microbial functional groups recovered in fertilized soils during the winter and nitrogen storage in microbial biomass was higher in fertilized soils in the autumn and winter than in the summer. Microbial immobilization of N may therefore be a significant sink for added N during autumn and winter months when plants are not active. One large microbial group that did not recover in the winter in fertilized soils was phenol mineralizers, possibly indicating selection against microbes with enzyme systems for the breakdown of phenolic compounds and complex soil organic matter. Overall, this work is a step towards understanding how chronic N additions affect the structure and biogeochemical functioning of soil microbial communities.     

Interannual variations of soil organic carbon fractions in unmanaged volcanic soils (Canary Islands,Spain)

The stability over time of the organic C stocked in soils under undisturbed ecosystems is poorly studied, despite being suitable for detecting changes related to climate fluctuations and global warming. Volcanic soils often show high organic C contents due to the stabilization of organic matter by short‐range ordered minerals or Al‐humus complexes. We investigated the dynamics of different organic C fractions in volcanic soils of protected natural ecosystems of the Canary Islands (Spain) to evaluate the stability of their C pools. The study was carried out in 10 plots, including both undisturbed and formerly disturbed ecosystems, over two annual periods. C inputs to (litterfall) and outputs from (respiration) the soil, root C stocks (0–30 cm), soil organic C (SOC) fractions belonging to C pools with different degrees of biogeochemical stability –total oxidisable C (TOC), microbial biomass C (MBC), water soluble C (WSC), hot‐water extractable C (HWC), humic C (HSC), – and total soil N (TN) (at 0–15 and 15–30 cm) were measured seasonally.A statistically significant interannual increase in CO₂ emissions and a decrease in the SOC, mainly at the expense of the most labile organic forms, were observed, while the root C stocks and litterfall inputs remained relatively constant over the study period. The observed changes may reflect an initial increase in SOC resulting from low soil respiration rates due to drought during the first year of study. The soils of nearly mature ecosystems were more apparently affected by C losses, while those undergoing the process of active natural regeneration exhibited disguised C loss because of the C sequestration trend that is characteristic of progressive ecological succession.     

Soil Organic Carbon Content Decreases in Both Surface and Subsoil Mineral Horizons by Simulated Future Increases in Labile Carbon Inputs in a Temperate Coniferous Forest

Soils represent important pools of soil organic carbon (SOC) that can be greatly influenced by labile C inputs, which are expected to increase in future due to CO₂ enrichment of atmosphere and a concomitant rise in plant primary productivity. Studying effects of variable labile C inputs on SOC pool helps to understand how soils respond to global change. However, this knowledge is missing for coniferous forest soils despite being widespread throughout the northern temperate zone. We conducted a 7-month field manipulation experiment to study the effects of variable labile C inputs (simulated by additions of C₄ sucrose) on the C content in soil fractions and on microbial abundance in the organic (O), surface mineral (A), and subsoil mineral (B) horizons of a temperate coniferous forest soil. SOC in less-protected soil fractions and total organic C were substantially decreased by labile C additions that simulated future increases in C inputs. The SOC losses were comparable between the A and B horizon (40% vs. 30%). However, because sucrose availability estimated from its incorporation into soil fractions and microbial biomass sharply decreased with soil depth, the loss of C was higher in the B than in the A horizon when related to the amount of sucrose added. Utilization of sucrose was highest by fungi in the O horizon and by bacteria in the mineral soil horizons. The results indicate that future increases in labile C inputs to coniferous forest soils will cause rapid and substantial losses of SOC in both the surface and subsoil mineral horizons.

     

Soil organic matter molecular composition with long-term detrital alterations is controlled by site-specific forest properties

Forest ecosystems are important global soil carbon (C) reservoirs, but their capacity to sequester C is susceptible to climate change factors that alter the quantity and quality of C inputs. To better understand forest soil C responses to altered C inputs, we integrated three molecular composition published data sets of soil organic matter (SOM) and soil microbial communities for mineral soils after 20 years of detrital input and removal treatments in two deciduous forests: Bousson Forest (BF), Harvard Forest (HF), and a coniferous forest: H.J. Andrews Forest (HJA). Soil C turnover times were estimated from radiocarbon measurements and compared with the molecular-level data (based on nuclear magnetic resonance and specific analysis of plant- and microbial-derived compounds) to better understand how ecosystem properties control soil C biogeochemistry and dynamics. Doubled aboveground litter additions did not increase soil C for any of the forests studied likely due to long-term soil priming. The degree of SOM decomposition was higher for bacteria-dominated sites with higher nitrogen (N) availability while lower for the N-poor coniferous forest. Litter exclusions significantly decreased soil C, increased SOM decomposition state, and led to the adaptation of the microbial communities to changes in available substrates. Finally, although aboveground litter determined soil C dynamics and its molecular composition in the coniferous forest (HJA), belowground litter appeared to be more influential in broadleaf deciduous forests (BH and HF). This synthesis demonstrates that inherent ecosystem properties regulate how soil C dynamics change with litter manipulations at the molecular-level. Across the forests studied, 20 years of litter additions did not enhance soil C content, whereas litter reductions negatively impacted soil C concentrations. These results indicate that soil C biogeochemistry at these temperate forests is highly sensitive to changes in litter deposition, which are a product of environmental change drivers.     

Grass invasion of a hardwood forest is associated with declines in belowground carbon pools

Invasive plant species affect a range of ecosystem processes but their impact on belowground carbon (C) pools is relatively unexplored. This is particularly true for grass invasions of forested ecosystems. Such invasions may alter both the quantity and quality of forest floor inputs. Dependent on both, two theories, ‘priming’ and ‘preferential substrate utilization’, suggest these changes may decrease, increase, or leave unchanged native plant‐derived soil C. Decreases are expected under ‘priming’ theory due to increased soil microbial activity. Under ‘preferential substrate utilization’, either an increase or no change is expected because the invasive plant's inputs are used by the microbial community instead of soil C. Here, we examine how Microstegium vimineum affects belowground C‐cycling in a southeastern US forest. Following predictions of priming theory, M. vimineum's presence is associated with decreases in native‐derived, C pools. For example, in September 2006 M. vimineum is associated with 24%, 34%, 36%, and 72% declines in total organic, particulate organic matter, mineralizable (a measure of microbially‐available C), and microbial biomass C, respectively. Soil C derived from M. vimineum does not compensate for these decreases, meaning that the sum of native‐ plus invasive‐derived C pools is smaller than native‐derived pools in uninvaded plots. Supporting our inferences that C‐cycling accelerates under invasion, the microbial community is more active per unit biomass: added ¹³C‐glucose is respired more rapidly in invaded plots. Our work suggests that this invader may accelerate C‐cycling in forest soils and deplete C stocks. The paucity of studies investigating impacts of grass invasion on C‐cycling in forests highlights the need to study further M. vimineum and other invasive grasses to assess their impacts on C sink strength and forest fertility.     

Increased microbial growth,biomass, and turnover drive soil organic carbon accumulation at higher plant diversity

Species‐rich plant communities have been shown to be more productive and to exhibit increased long‐term soil organic carbon (SOC) storage. Soil microorganisms are central to the conversion of plant organic matter into SOC, yet the relationship between plant diversity, soil microbial growth, turnover as well as carbon use efficiency (CUE) and SOC accumulation is unknown. As heterotrophic soil microbes are primarily carbon limited, it is important to understand how they respond to increased plant‐derived carbon inputs at higher plant species richness (PSR). We used the long‐term grassland biodiversity experiment in Jena, Germany, to examine how microbial physiology responds to changes in plant diversity and how this affects SOC content. The Jena Experiment considers different numbers of species (1–60), functional groups (1–4) as well as functional identity (small herbs, tall herbs, grasses, and legumes). We found that PSR accelerated microbial growth and turnover and increased microbial biomass and necromass. PSR also accelerated microbial respiration, but this effect was less strong than for microbial growth. In contrast, PSR did not affect microbial CUE or biomass‐specific respiration. Structural equation models revealed that PSR had direct positive effects on root biomass, and thereby on microbial growth and microbial biomass carbon. Finally, PSR increased SOC content via its positive influence on microbial biomass carbon. We suggest that PSR favors faster rates of microbial growth and turnover, likely due to greater plant productivity, resulting in higher amounts of microbial biomass and necromass that translate into the observed increase in SOC. We thus identify the microbial mechanism linking species‐rich plant communities to a carbon cycle process of importance to Earth's climate system.     

Indirect Effects of Nitrogen Amendments on Organic Substrate Quality Increase Enzymatic Activity Driving Decomposition in a Mesic Grassland

The fate of soil organic carbon (SOC) is determined, in part, by complex interactions between the quality of plant litter inputs, nutrient availability, and the microbial communities that control decomposition rates. This study explores these interactions in a mesic grassland where C and nitrogen (N) availability and plant litter quality have been manipulated using both fertilization and haying for 7 years. We measured a suite of soil parameters including inorganic N, extractable organic C and N (EOC and EON), soil moisture, extracellular enzyme activity (EEA), and the isotopic composition of C and N in the microbial biomass and substrate sources. We use these data to determine how the activity of microbial decomposers was influenced by varying levels of substrate C and N quality and quantity and to explore potential mechanisms explaining the fate of enhanced plant biomass inputs with fertilization. Oxidative EEA targeting relatively recalcitrant C pools was not affected by fertilization. EEA linked to the breakdown of relatively labile C rich substrates exhibited no relationship with inorganic N availability but was significantly greater with fertilization and associated increases in substrate quality. These increases in EEA were not related to an increase in microbial biomass C. The ratio of hydrolytic C:N acquisition enzymes and δ¹³C and δ¹⁵N values of microbial biomass relative to bulk soil C and N, or EOC and EON suggest that microbial communities in fertilized plots were relatively C limited, a feature likely driving enhanced microbial efforts to acquire C from labile sources. These data suggest that in mesic grasslands, enhancements in biomass inputs and quality with fertilization can prompt an increase in EEA within the mineral soil profile with no significant increases in microbial biomass. Our work helps elucidate the microbially mediated fate of enhanced biomass inputs that are greater in magnitude than the associated increases in mineral soil organic matter.     

Consistent effects of nitrogen amendments on soil microbial communities and processes across biomes

Ecosystems worldwide are receiving increasing amounts of reactive nitrogen (N) via anthropogenic activities with the added N having potentially important impacts on microbially mediated belowground carbon dynamics. However, a comprehensive understanding of how elevated N availability affects soil microbial processes and community dynamics remains incomplete. The mechanisms responsible for the observed responses are poorly resolved and we do not know if soil microbial communities respond in a similar manner across ecosystems. We collected 28 soils from a broad range of ecosystems in North America, amended soils with inorganic N, and incubated the soils under controlled conditions for 1 year. Consistent across nearly all soils, N addition decreased microbial respiration rates, with an average decrease of 11% over the year‐long incubation, and decreased microbial biomass by 35%. High‐throughput pyrosequencing showed that N addition consistently altered bacterial community composition, increasing the relative abundance of Actinobacteria and Firmicutes, and decreasing the relative abundance of Acidobacteria and Verrucomicrobia. Further, N‐amended soils consistently had lower activities in a broad suite of extracellular enzymes and had decreased temperature sensitivity, suggesting a shift to the preferential decomposition of more labile C pools. The observed trends held across strong gradients in climate and soil characteristics, indicating that the soil microbial responses to N addition are likely controlled by similar wide‐spread mechanisms. Our results support the hypothesis that N addition depresses soil microbial activity by shifting the metabolic capabilities of soil bacterial communities, yielding communities that are less capable of decomposing more recalcitrant soil carbon pools and leading to a potential increase in soil carbon sequestration rates.     

贵州山区土壤中微生物担是能源物质碳流动的源与汇

在传统的农业生态系统的研究中 ,主要精力放在营养物 (如Ｎ)上 ,认为它们是限制生产力的因素 ;而往往忽略了土壤中碳的重要性 ,认为收获不受Ｃ限制的影响。然而 ,碳循环中的有机碳的分解作用部分控制着出现在地表下和显露在地表上的农业过程[4]。土壤中所储存的有机质 ,其数量既反映土壤从植物残留物的输入所获得的有机质与微生物群落的能量和营养需求之间的平衡 ,又反映植物对营养物的需求与有机质分解作用之间的平衡。因此 ,土壤中碳的平衡能反映出有机质中能量物质的储存[5]。大部分由光合作用形成的碳 ,是通过地表下的生态系统来流动的[…     

Improved model simulation of soil carbon cycling by representing the microbially derived organic carbon pool

During the decomposition process of soil organic carbon (SOC), microbial products such as microbial necromass and microbial metabolites may form an important stable carbon (C) pool, called microbially derived C, which has different decomposition patterns from plant-derived C. However, current Earth System Models do not simulate this microbially derived C pool separately. Here, we incorporated the microbial necromass pool to the first-order kinetic model and the Michaelis–Menten model, respectively, and validated model behaviors against previous observation data from the decomposition experiments of ¹³C-labeled necromass. Our models showed better performance than existing models and the Michaelis–Menten model was better than the first-order kinetic model. Microbial necromass C was estimated to be 10–27% of total SOC in the study soils by our models and therefore should not be ignored. This study provides a novel modification to process-based models for better simulation of soil organic C under the context of global changes.Subject terms: Biogeochemistry, Theoretical ecology, Microbial ecology, Stable isotope analysis     

Climate change effects on the stability and chemistry of soil organic carbon pools in a subalpine grassland

Mountain soils stock large quantities of carbon as particulate organic matter that may be highly vulnerable to climate change. To explore potential shifts in soil organic matter (SOM) form and stability under climate change (warming and reduced precipitations), we studied the dynamics of SOM pools of a mountain grassland in the Swiss Jura as part of a climate manipulation experiment. The climate manipulation (elevational soil transplantation) was set up in October 2009 and simulated two realistic climate change scenarios. After 4 years of manipulation, we performed SOM physical fractionation to extract SOM fractions corresponding to specific turnover rates, in winter and in summer. Soil organic matter fraction chemistry was studied with ultraviolet, 3D fluorescence, and mid-infrared spectroscopies. The most labile SOM fractions showed high intra-annual dynamics (amounts and chemistry) mediated via the seasonal changes of fresh plant debris inputs and confirming their high contribution to the microbial loop. Our climate change manipulation modified the chemical differences between free and intra-aggregate organic matter, suggesting a modification of soil macro-aggregates dynamics. Interestingly, the 4-year climate manipulation affected directly the SOM dynamics, with a decrease in organic C bulk soil content, resulting from significant C-losses in the mineral-associated SOM fraction (MAOM), the most stable form of SOM. This SOC decrease was associated with a decrease in clay content, above- and belowground plants biomass, soil microbial biomass and activity. The combination of these climate changes effects on the plant–soil system could have led to increase C-losses from the MAOM fraction through clay-SOM washing out and DOC leaching in this subalpine grassland.     

Introduced ungulate herbivore alters soil processes after fire

Ungulate herbivory can have profound effects on ecosystem processes by altering organic inputs of leaves and roots as well as changing soil physical and chemical properties. These effects may be especially important when the herbivore is an introduced species. Utilizing large mammal exclosures to prevent access by introduced elk at multiple sites along a fire chronosequence, we examined the effects of elk herbivory and fire on soil microbial activity and nutrient availability. Using time since fire as a co-variate and herbivore exclosures, paired with areas outside of the exclosures, we hypothesized that reductions in plant biomass due to herbivory would reduce organic inputs to soils and impact soil microbial activities and nutrient storage. We found three major patterns: (1) when elk were excluded, surface mineral soils had higher soil organic carbon (C), total nitrogen (N), microbial N pools, and increased extra-cellular enzyme activity of a C-acquiring enzyme across a gradient of time since fire. (2) When introduced elk are present, the activity of some extracellular enzymes as well as NO₃ ⁻ availability are enhanced in the soil but the post-fire patterns described above with respect to nutrient accrual over time are delayed. (3) Herbivory by an introduced ungulate upsets the trajectory of ecosystem “recovery” after wildfire and delays soil C and N dynamics by an estimated 14.5–21 years, respectively. These results suggest that introduced, browsing herbivores significantly decelerate ecosystem processes but herbivory by exotics may also result in unpredictability in specific soil responses.     

Grazing enhances belowground carbon allocation,microbial biomass,and soil carbon in a subtropical grassland

Despite the large contribution of rangeland and pasture to global soil organic carbon (SOC) stocks, there is considerable uncertainty about the impact of large herbivore grazing on SOC, especially for understudied subtropical grazing lands. It is well known that root system inputs are the source of most grassland SOC, but the impact of grazing on partitioning of carbon allocation to root tissue production compared to fine root exudation is unclear. Given that different forms of root C have differing implications for SOC synthesis and decomposition, this represents a significant gap in knowledge. Root exudates should contribute to SOC primarily after microbial assimilation, and thus promote microbial contributions to SOC based on stabilization of microbial necromass, whereas root litter deposition contributes directly as plant‐derived SOC following microbial decomposition. Here, we used in situ isotope pulse‐chase methodology paired with plant and soil sampling to link plant carbon allocation patterns with SOC pools in replicated long‐term grazing exclosures in subtropical pasture in Florida, USA. We quantified allocation of carbon to root tissue and measured root exudation across grazed and ungrazed plots and quantified lignin phenols to assess the relative contribution of microbial vs. plant products to total SOC. We found that grazing exclusion was associated with dramatically less overall belowground allocation, with lower root biomass, fine root exudates, and microbial biomass. Concurrently, grazed pasture contained greater total SOC, and a larger fraction of SOC that originated from plant tissue deposition, suggesting that higher root litter deposition under grazing promotes greater SOC. We conclude that grazing effects on SOC depend on root system biomass, a pattern that may generalize to other C4‐dominated grasslands, especially in the subtropics. Improved understanding of ecological factors underlying root system biomass may be the key to forecasting SOC and optimizing grazing management to enhance SOC accumulation.