Optimal metabolic regulation along resource stoichiometry gradients

Most heterotrophic organisms feed on substrates that are poor in nutrients compared to their demand, leading to elemental imbalances that may constrain their growth and function. Flexible carbon (C)‐use efficiency (CUE, C used for growth over C taken up) can represent a strategy to reduce elemental imbalances. Here, we argue that metabolic regulation has evolved to maximise the organism growth rate along gradients of nutrient availability and translated this assumption into an optimality model that links CUE to substrate and organism stoichiometry. The optimal CUE is predicted to decrease with increasing substrate C‐to‐nutrient ratio, and increase with nutrient amendment. These predictions are generally confirmed by empirical evidence from a new database of c. 2200 CUE estimates, lending support to the hypothesis that CUE is optimised across levels of organisation (microorganisms and animals), in aquatic and terrestrial systems, and when considering nitrogen or phosphorus as limiting nutrients.     

Carbon use efficiency of microbial communities: stoichiometry,methodology and modelling

Carbon use efficiency (CUE) is a fundamental parameter for ecological models based on the physiology of microorganisms. CUE determines energy and material flows to higher trophic levels, conversion of plant‐produced carbon into microbial products and rates of ecosystem carbon storage. Thermodynamic calculations support a maximum CUE value of ~ 0.60 (CUE _max). Kinetic and stoichiometric constraints on microbial growth suggest that CUE in multi‐resource limited natural systems should approach ~ 0.3 (CUE _max/2). However, the mean CUE values reported for aquatic and terrestrial ecosystems differ by twofold (~ 0.26 vs. ~ 0.55) because the methods used to estimate CUE in aquatic and terrestrial systems generally differ and soil estimates are less likely to capture the full maintenance costs of community metabolism given the difficulty of measurements in water‐limited environments. Moreover, many simulation models lack adequate representation of energy spilling pathways and stoichiometric constraints on metabolism, which can also lead to overestimates of CUE. We recommend that broad‐scale models use a CUE value of 0.30, unless there is evidence for lower values as a result of pervasive nutrient limitations. Ecosystem models operating at finer scales should consider resource composition, stoichiometric constraints and biomass composition, as well as environmental drivers, to predict the CUE of microbial communities.     

Microbial carbon use efficiency in different ecosystems: A meta-analysis based on a biogeochemical equilibrium model

Soil microbial carbon use efficiency (CUE) is a crucial parameter that can be used to evaluate the partitioning of soil carbon (C) between microbial growth and respiration. However, general patterns of microbial CUE among terrestrial ecosystems (e.g., farmland, grassland, and forest) remain controversial. To address this knowledge gap, data from 41 study sites (n = 197 soil samples) including 58 farmlands, 95 forests, and 44 grasslands were collected and analyzed to estimate microbial CUEs using a biogeochemical equilibrium model. We also evaluated the metabolic limitations of microbial growth using an enzyme vector model and the drivers of CUE across different ecosystems. The CUEs obtained from soils of farmland, forest, and grassland ecosystems were significantly different with means of 0.39, 0.33, and 0.42, respectively, illustrating that grassland soils exhibited higher microbial C sequestration potentials (p < .05). Microbial metabolic limitations were also distinct in these ecosystems, and carbon limitation was dominant exhibiting strong negative effects on CUE. Exoenzyme stoichiometry played a greater role in impacting CUE values than soil elemental stoichiometry within each ecosystem. Specifically, soil exoenzymatic ratios of C:phosphorus (P) acquisition activities (EEA_C:P) and the exoenzymatic ratio of C:nitrogen (N) acquisition activities (EEA_C:N) imparted strong negative effects on soil microbial CUE in grassland and forest ecosystems, respectively. But in farmland soils, EEA_C:P exhibited greater positive effects, showing that resource constraints could regulate microbial resource allocation with discriminating patterns across terrestrial ecosystems. Furthermore, mean annual temperature (MAT) rather than mean annual precipitation (MAP) was a critical climate factor affecting CUE, and soil pH as a major factor remained positive to drive the changes in microbial CUE within ecosystems. This research illustrates a conceptual framework of microbial CUEs in terrestrial ecosystems and provides the theoretical evidence to improve soil microbial C sequestration capacity in response to global change.     

Increasing microbial carbon use efficiency with warming predicts soil heterotrophic respiration globally

The degree to which climate warming will stimulate soil organic carbon (SOC) losses via heterotrophic respiration remains uncertain, in part because different or even opposite microbial physiology and temperature relationships have been proposed in SOC models. We incorporated competing microbial carbon use efficiency (CUE)–mean annual temperature (MAT) and enzyme kinetic–MAT relationships into SOC models, and compared the simulated mass‐specific soil heterotrophic respiration rates with multiple published datasets of measured respiration. The measured data included 110 dryland soils globally distributed and two continental to global‐scale cross‐biome datasets. Model–data comparisons suggested that a positive CUE–MAT relationship best predicts the measured mass‐specific soil heterotrophic respiration rates in soils distributed globally. These results are robust when considering models of increasing complexity and competing mechanisms driving soil heterotrophic respiration–MAT relationships (e.g., carbon substrate availability). Our findings suggest that a warmer climate selects for microbial communities with higher CUE, as opposed to the often hypothesized reductions in CUE by warming based on soil laboratory assays. Our results help to build the impetus for, and confidence in, including microbial mechanisms in soil biogeochemical models used to forecast changes in global soil carbon stocks in response to warming.     

Microbial Growth and Carbon Use Efficiency in the Rhizosphere and Root-Free Soil

Plant-microbial interactions alter C and N balance in the rhizosphere and affect the microbial carbon use efficiency (CUE)–the fundamental characteristic of microbial metabolism. Estimation of CUE in microbial hotspots with high dynamics of activity and changes of microbial physiological state from dormancy to activity is a challenge in soil microbiology. We analyzed respiratory activity, microbial DNA content and CUE by manipulation the C and nutrients availability in the soil under Beta vulgaris. All measurements were done in root-free and rhizosphere soil under steady-state conditions and during microbial growth induced by addition of glucose. Microorganisms in the rhizosphere and root-free soil differed in their CUE dynamics due to varying time delays between respiration burst and DNA increase. Constant CUE in an exponentially-growing microbial community in rhizosphere demonstrated the balanced growth. In contrast, the CUE in the root-free soil increased more than three times at the end of exponential growth and was 1.5 times higher than in the rhizosphere. Plants alter the dynamics of microbial CUE by balancing the catabolic and anabolic processes, which were decoupled in the root-free soil. The effects of N and C availability on CUE in rhizosphere and root-free soil are discussed.     

Ecoenzymatic stoichiometry of stream sediments with comparison to terrestrial soils

The kinetics and elemental composition of cellular units that mediate production and respiration are the basis for the metabolic and stoichiometric theories of ecological organization. This theoretical framework extends to the activities of microbial enzymes released into the environment (ecoenzymes) that mediate the release of assimilable substrate from detrital organic matter. In this paper, we analyze the stoichiometry of ecoenzymatic activities in the surface sediments of lotic ecosystems and compare those results to the stoichiometry observed in terrestrial soils. We relate these ecoenzymatic ratios to energy and nutrient availability in the environment as well as microbial elemental content and growth efficiency. The data, collected by US Environmental Protection Agency, include the potential activities of 11 enzymes for 2,200 samples collected across the US, along with analyses of sediment C, N and P content. On average, ecoenzymatic activities in stream sediments are 2–5 times greater per gC than those of terrestrial soils. Ecoenzymatic ratios of C, N and P acquisition activities support elemental analyses showing that microbial metabolism is more likely to be C-limited than N or P-limited compared to terrestrial soils. Ratios of hydrolytic to oxidative activities indicate that sediment organic matter is more labile than soil organic matter and N acquisition is less dependent on humic oxidation. The mean activity ratios of glycosidases and aminopeptidases reflect the environmental abundance of their respective substrates. For both freshwater sediments and terrestrial soils, the mean C:nutrient ratio of microbial biomass normalized to growth efficiency approximates the mean ecoenzymatic C:nutrient activity ratios normalized to environmental C:nutrient abundance. This relationship defines a condition for biogeochemical equilibrium consistent with stoichiometric and metabolic theory.     

Microbial community dynamics alleviate stoichiometric constraints during litter decay

Under the current paradigm, organic matter decomposition and nutrient cycling rates are a function of the imbalance between substrate and microbial biomass stoichiometry. Challenging this view, we demonstrate that in an individual‐based model, microbial community dynamics alter relative C and N limitation during litter decomposition, leading to a system behaviour not predictable from stoichiometric theory alone. Rather, the dynamics of interacting functional groups lead to an adaptation at the community level, which accelerates nitrogen recycling in litter with high initial C : N ratios and thus alleviates microbial N limitation. This mechanism allows microbial decomposers to overcome large imbalances between resource and biomass stoichiometry without the need to decrease carbon use efficiency (CUE), which is in contrast to predictions of traditional stoichiometric mass balance equations. We conclude that identifying and implementing microbial community‐driven mechanisms in biogeochemical models are necessary for accurately predicting terrestrial C fluxes in response to changing environmental conditions.     

Inter-annual Variability of Soil Respiration in Wet Shrublands: Do Plants Modulate Its Sensitivity to Climate?

Understanding the response of soil respiration to climate variability is critical to formulate realistic predictions of future carbon (C) fluxes under different climate change scenarios. There is growing evidence that the influence of long-term climate variability in C fluxes from terrestrial ecosystems is modulated by adjustments in the aboveground–belowground links. Here, we studied the inter-annual variability in soil respiration from a wet shrubland going through successional change in North Wales (UK) during 13 years. We hypothesised that the decline in plant productivity observed over a decade would result in a decrease in the apparent sensitivity of soil respiration to soil temperature, and that rainfall variability would explain a significant fraction of the inter-annual variability in plant productivity, and consequently, in soil respiration, due to excess-water constraining nutrient availability for plants. As hypothesised, there were parallel decreases between plant productivity and annual and summer CO₂ emissions over the 13-year period. Soil temperatures did not follow a similar trend, which resulted in a decline in the apparent sensitivity of soil respiration to soil temperature (apparent Q₁₀ values decreased from 9.4 to 2.8). Contrary to our second hypothesis, summer maximum air temperature rather than rainfall was the climate variable with the greatest influence on aboveground biomass and annual cumulative respiration. Since summer air temperature and rainfall were positively associated, the greatest annual respiration values were recorded during years of high rainfall. The results suggest that adjustments in plant productivity might have a critical role in determining the long-term-sensitivity of soil respiration to changing climate conditions.     

Ecoenzymatic stoichiometry and microbial processing of organic matter in northern bogs and fens reveals a common P-limitation between peatland types

We compared carbon (C), nitrogen (N), and phosphorus (P) concentrations in atmospheric deposition, runoff, and soils with microbial respiration [dehydrogenase (DHA)] and ecoenzyme activity (EEA) in an ombrotrophic bog and a minerotrophic fen to investigate the environmental drivers of biogeochemical cycling in peatlands at the Marcell Experimental Forest in northern Minnesota, USA. Ecoenzymatic stoichiometry was used to construct models for C use efficiency (CUE) and decomposition (M), and these were used to model respiration (R_m). Our goals were to determine the relative C, N, and P limitations on microbial processes and organic matter decomposition, and to identify environmental constraints on ecoenzymatic processes. Mean annual water, C, and P yields were greater in the fen, while N yields were similar in both the bog and fen. Soil chemistry differed between the bog and fen, and both watersheds exhibited significant differences among soil horizons. DHA and EEA differed by watersheds and soil horizons, CUE, M, and R_m differed only by soil horizons. C, N, or P limitations indicated by EEA stoichiometry were confirmed with orthogonal regressions of ecoenzyme pairs and enzyme vector analyses, and indicated greater N and P limitation in the bog than in the fen, with an overall tendency toward P-limitation in both the bog and fen. Ecoenzymatic stoichiometry, microbial respiration, and organic matter decomposition were responsive to resource availability and the environmental drivers of microbial metabolism, including those related to global climate changes.     

A Framework to Assess Biogeochemical Response to Ecosystem Disturbance Using Nutrient Partitioning Ratios

Disturbances affect almost all terrestrial ecosystems, but it has been difficult to identify general principles regarding these influences. To improve our understanding of the long-term consequences of disturbance on terrestrial ecosystems, we present a conceptual framework that analyzes disturbances by their biogeochemical impacts. We posit that the ratio of soil and plant nutrient stocks in mature ecosystems represents a characteristic site property. Focusing on nitrogen (N), we hypothesize that this partitioning ratio (soil N: plant N) will undergo a predictable trajectory after disturbance. We investigate the nature of this partitioning ratio with three approaches: (1) nutrient stock data from forested ecosystems in North America, (2) a process-based ecosystem model, and (3) conceptual shifts in site nutrient availability with altered disturbance frequency. Partitioning ratios could be applied to a variety of ecosystems and successional states, allowing for improved temporal scaling of disturbance events. The generally short-term empirical evidence for recovery trajectories of nutrient stocks and partitioning ratios suggests two areas for future research. First, we need to recognize and quantify how disturbance effects can be accreting or depleting, depending on whether their net effect is to increase or decrease ecosystem nutrient stocks. Second, we need to test how altered disturbance frequencies from the present state may be constructive or destructive in their effects on biogeochemical cycling and nutrient availability. Long-term studies, with repeated sampling of soils and vegetation, will be essential in further developing this framework of biogeochemical response to disturbance.     

Microbial ecoenzyme stoichiometry,nutrient limitation,and organic matter decomposition in wetlands of the conterminous United States

Microbial respiration (R_m) and ecoenzyme activities (EEA) related to microbial carbon, nitrogen, and phosphorus acquisition were measured in 792 freshwater and estuarine wetlands (representing a cumulative area of 217,480 km²) across the continental United States as part of the US EPA’s 2011 National Wetland Condition Assessment. EEA stoichiometry was used to construct models for and assess nutrient limitation, carbon use efficiency (CUE), and organic matter decomposition (? k). The wetlands were classified into ten groups based on aggregated ecoregion and wetland type. The wetlands were also assigned to least, intermediate, and most disturbed classes, based on the extent of human influences. Ecoenzyme activity related to C, N and P acquisition, R_m, CUE, and ? k differed among ecoregion–wetland types and, with the exception of C acquisition and ? k, among disturbance classes. R_m and EEA were positively correlated with soil C, N and P content (r = 0.15–0.64) and stoichiometry (r = 0.15–0.48), and negatively correlated with an index of carbon quality (r = ? 0.22 to ? 0.39). EEA stoichiometry revealed that wetlands were more often P- than N-limited, and that P-limitation increases with increasing disturbance. Our enzyme-based approach for modeling C, N, and P acquisition, and organic matter decomposition, all rooted in stoichiometric theory, provides a mechanism for modeling resource limitations of microbial metabolism and biogeochemical cycling in wetlands. Given the ease of collecting and analyzing soil EEA and their response to wetland disturbance gradients, enzyme stoichiometry models are a cost-effective tool for monitoring ecosystem responses to resource availability and the environmental drivers of microbial metabolism, including those related to global climate changes.     

Fungal interactions reduce carbon use efficiency

The efficiency by which fungi decompose organic matter contributes to the amount of carbon that is retained in biomass vs. lost to the atmosphere as respiration. This carbon use efficiency (CUE) is affected by various abiotic conditions, including temperature and nutrient availability. Theoretically, the physiological costs of interspecific interactions should likewise alter CUE, yet the magnitude of these costs is untested. Here we conduct a microcosm experiment to quantify how interactions among wood‐decay basidiomycete fungi alter growth, respiration and CUE across a temperature and nitrogen gradient. We show that species interactions induced consistent declines in CUE, regardless of abiotic conditions. Multispecies communities exhibited reductions in CUE of up to 25% relative to individual CUE, with this biotic effect being greater than the observed variation attributable to abiotic conditions. Our results suggest that the extent to which fungal‐mediated carbon fluxes respond to environmental change may be influenced strongly by species interactions.     

土壤微生物碳素利用效率研究进展

土壤微生物碳素利用效率(CUE)是指微生物将吸收的碳(C)转化为自身生物量C的效率,也称为微生物的生长效率。土壤微生物CUE是生态系统C循环中的重要生理生态学参数,影响着生态系统的C固持、周转、土壤矿化以及温室气体排放等过程。在全球环境变化背景下,认识土壤微生物CUE的变异及其影响机制,对于更好的认识生态系统C循环过程及其对全球变化的响应具有重要意义。概述了CUE的定义及其测定方法,重点综述和分析土壤微生物CUE的变异及影响因素取得的研究进展。基于现有研究的分析得出,土壤微生物CUE通常表示为微生物的生长与吸收的比值,分为基于微生物生长速率、微生物生物量、底物吸收速率和底物浓度变化等方法进行测定。土壤微生物CUE在0.2—0.8的范围内变化,这种变异主要受到来自热力学、生态环境因子、底物养分质量和有效性、化学计量平衡以及微生物群落组成的影响。今后土壤微生物CUE的研究应加强对微量代谢组分的定量分析,生物和环境要素交互影响的调控机理解析,以及微生物动态生理响应过程的碳循环模型优化。     

Stoichiometric tracking of soil nutrients by a desert insect herbivore

Abstract Biogeochemistry and population biology have developed independently, with few attempts at linkage, almost none of which were mechanistically based. We hypothesize that biogeochemical cycling is linked to herbivore population dynamics through the influence of soil nutrient availability on foliar nutrient content, which constrains herbivore investment in phosphorus (P)‐rich molecules necessary for growth. We show that variation in desert soil P availability is linked to abundance of an insect herbivore (Sabinia setosa) through the influence of soil P on the C:P ratio of the host plant (Prosopis velutina). Low P availability increases C:P ratio of Prosopis leaves, leading to a decline in body %P, %RNA and abundance of Sabinia. Tight association between soil, plant and herbivore P provides the first evidence of a mechanistic pathway linking P biogeochemistry to terrestrial food webs by altering the supply of dietary P to herbivores, thus limiting their capacity for growth by constraining the production of P‐rich cellular ribosomal RNA (rRNA).     

Does declining carbon‐use efficiency explain thermal acclimation of soil respiration with warming?

Enhanced soil respiration in response to global warming may substantially increase atmospheric CO₂ concentrations above the anthropogenic contribution, depending on the mechanisms underlying the temperature sensitivity of soil respiration. Here, we compared short‐term and seasonal responses of soil respiration to a shifting thermal environment and variable substrate availability via laboratory incubations. To analyze the data from incubations, we implemented a novel process‐based model of soil respiration in a hierarchical Bayesian framework. Our process model combined a Michaelis–Menten‐type equation of substrate availability and microbial biomass with an Arrhenius‐type nonlinear temperature response function. We tested the competing hypotheses that apparent thermal acclimation of soil respiration can be explained by depletion of labile substrates in warmed soils, or that physiological acclimation reduces respiration rates. We demonstrated that short‐term apparent acclimation can be induced by substrate depletion, but that decreasing microbial biomass carbon (MBC) is also important, and lower MBC at warmer temperatures is likely due to decreased carbon‐use efficiency (CUE). Observed seasonal acclimation of soil respiration was associated with higher CUE and lower basal respiration for summer‐ vs. winter‐collected soils. Whether the observed short‐term decrease in CUE or the seasonal acclimation of CUE with increased temperatures dominates the response to long‐term warming will have important consequences for soil organic carbon storage.     

Cascading effects of induced terrestrial plant defences on aquatic and terrestrial ecosystem function

Herbivores induce plants to undergo diverse processes that minimize costs to the plant, such as producing defences to deter herbivory or reallocating limited resources to inaccessible portions of the plant. Yet most plant tissue is consumed by decomposers, not herbivores, and these defensive processes aimed to deter herbivores may alter plant tissue even after detachment from the plant. All consumers value nutrients, but plants also require these nutrients for primary functions and defensive processes. We experimentally simulated herbivory with and without nutrient additions on red alder (Alnus rubra), which supplies the majority of leaf litter for many rivers in western North America. Simulated herbivory induced a defence response with cascading effects: terrestrial herbivores and aquatic decomposers fed less on leaves from stressed trees. This effect was context dependent: leaves from fertilized-only trees decomposed most rapidly while leaves from fertilized trees receiving the herbivory treatment decomposed least, suggesting plants funnelled a nutritionally valuable resource into enhanced defence. One component of the defence response was a decrease in leaf nitrogen leading to elevated carbon : nitrogen. Aquatic decomposers prefer leaves naturally low in C : N and this altered nutrient profile largely explains the lower rate of aquatic decomposition. Furthermore, terrestrial soil decomposers were unaffected by either treatment but did show a preference for local and nitrogen-rich leaves. Our study illustrates the ecological implications of terrestrial herbivory and these findings demonstrate that the effects of selection caused by terrestrial herbivory in one ecosystem can indirectly shape the structure of other ecosystems through ecological fluxes across boundaries.     

罗浮栲林土壤微生物碳利用效率对短期氮添加的响应

作为调节土壤碳矿化过程的重要参数,微生物碳利用效率(CUE)对理解陆地生态系统中的碳循环至关重要。本研究在戴云山罗浮栲林设置对照(0 kg N·hm^-2·a^-1)、低氮(40 kg N·hm^-2·a^-1)和高氮(80 kg N·hm^-2·a^-1) 3个氮添加水平以模拟氮沉降,测定了表层(0～10 cm)土壤基本理化性质、有机碳组分、微生物生物量和酶活性;并利用¹⁸O标记水方法测定土壤微生物CUE,以更好地理解氮沉降加剧对微生物CUE的影响及其影响因素。结果表明: 短期氮添加显著降低了土壤微生物的呼吸速率、碳和氮获取酶活性,但显著增加了土壤微生物CUE。β-N-乙酰氨基酸葡糖苷酶(NAG)/微生物生物量碳(MBC)、微生物呼吸速率、β-葡萄糖苷酶(BG)/MBC、纤维素水解酶(CBH)/MBC和土壤有机碳含量是影响CUE的主要因素,且CUE与NAG/MBC、微生物呼吸速率、BG/MBC和CBH/MBC呈显著负相关,与土壤有机碳呈显著正相关。综上,短期氮添加导致土壤微生物获取碳和氮的成本降低,减少微生物呼吸,从而提高了土壤微生物CUE,这将有助于提高罗浮栲林土壤碳固存潜力。     

What controls variation in carbon use efficiency among Amazonian tropical forests?

Why do some forests produce biomass more efficiently than others? Variations in Carbon Use Efficiency (CUE: total Net Primary Production (NPP)/ Gross Primary Production (GPP)) may be due to changes in wood residence time (Biomass/NPP_wood), temperature, or soil nutrient status. We tested these hypotheses in 14, one ha plots across Amazonian and Andean forests where we measured most key components of net primary production (NPP: wood, fine roots, and leaves) and autotrophic respiration (R_a; wood, rhizosphere, and leaf respiration). We found that lower fertility sites were less efficient at producing biomass and had higher rhizosphere respiration, indicating increased carbon allocation to belowground components. We then compared wood respiration to wood growth and rhizosphere respiration to fine root growth and found that forests with residence times <40 yrs had significantly lower maintenance respiration for both wood and fine roots than forests with residence times >40 yrs. A comparison of rhizosphere respiration to fine root growth showed that rhizosphere growth respiration was significantly greater at low fertility sites. Overall, we found that Amazonian forests produce biomass less efficiently in stands with residence times >40 yrs and in stands with lower fertility, but changes to long‐term mean annual temperatures do not impact CUE.     

Are patterns in nutrient limitation belowground consistent with those aboveground: results from a 4 million year chronosequence

Accurately predicting the effects of global change on net carbon (C) exchange between terrestrial ecosystems and the atmosphere requires a more complete understanding of how nutrient availability regulates both plant growth and heterotrophic soil respiration. Models of soil development suggest that the nature of nutrient limitation changes over the course of ecosystem development, transitioning from nitrogen (N) limitation in ‘young’ sites to phosphorus (P) limitation in ‘old’ sites. However, previous research has focused primarily on plant responses to added nutrients, and the applicability of nutrient limitation-soil development models to belowground processes has not been thoroughly investigated. Here, we assessed the effects of nutrients on soil C cycling in three different forests that occupy a 4 million year substrate age chronosequence where tree growth is N limited at the youngest site, co-limited by N and P at the intermediate-aged site, and P limited at the oldest site. Our goal was to use short-term laboratory soil C manipulations (using ¹⁴C-labeled substrates) and longer-term intact soil core incubations to compare belowground responses to fertilization with aboveground patterns. When nutrients were applied with labile C (sucrose), patterns of microbial nutrient limitation were similar to plant patterns: microbial activity was limited more by N than by P in the young site, and P was more limiting than N in the old site. However, in the absence of C additions, increased respiration of native soil organic matter only occurred with simultaneous additions of N and P. Taken together, these data suggest that altered nutrient inputs into ecosystems could have dissimilar effects on C cycling above- and belowground, that nutrients may differentially affect of the fate of different soil C pools, and that future changes to the net C balance of terrestrial ecosystems will be partially regulated by soil nutrient status.     

Evaluating soil microbial carbon use efficiency explicitly as a function of cellular processes: implications for measurements and models

Carbon use efficiency (CUE), the proportion of carbon (C) consumed by microbes that is converted into biomass, is an important parameter for soil C models with explicit microbial controls. While often considered as a single parameter, CUE is an emergent property of multiple microbial processes, including assimilation efficiency, biomass-specific respiration, enzyme production, and respiratory costs of enzyme production. These processes occur over variable time scales and imply different fates for C, and the same emergent CUE value can result when C is allocated in fundamentally different ways (e.g. a high investment in enzyme production vs. a high assimilation cost). We developed a model that represents the individual processes underlying emergent CUE to test how shifts in microbial allocation alter equilibrium soil C pool sizes. We found that an increase in emergent CUE that results from a change in assimilation efficiency, biomass specific respiration, or respiration costs from enzyme production causes soil organic C (SOC) to decline, while the same change in emergent CUE resulting from a change in enzyme production causes SOC to increase. We also used the model to test the sensitivity of CUE from isotopic C tracer estimates to changes in microbial allocation processes. We found that these estimates do not account for the same microbial processes represented by emergent CUE in models. We propose that considering microbial processes explicitly rather than representing CUE as a single parameter can improve data-model integration. In addition, modeling microbial processes explicitly will account for a wider range of possible outcomes from shifts in microbial C allocation, such as when increased SOC results from increasing CUE.