Microbial Biomass Stoichiometry Research Articles

C/P stoichiometry of dying rice root defines the spatial distribution and dynamics of enzyme activities in root-detritusphere

As the primary microbial substrate after shoot cutting, the element stoichiometry of root-detritus (dying or dead roots) influences the enzyme activity in root-detritusphere. However, the effect of the C/P ratio of root-detritus on the dynamics and distribution of enzyme activities is little revealed. We hypothesised that P fertilisation would decrease the C/P ratio of root-detritus, therefore affecting the hotspot areas and hot moments of C-acquiring and P-acquiring enzyme activities, as well as their activity ratio (C/P acquisition ratio). Root-detritus of low (59.0) and high (170.8) C/P ratios was produced in P-poor soil with and without P fertilisation, respectively. In situ soil zymography showed that the distribution of C-acquiring enzymes (β-glucosidase and cellobiohydrolase) was more associated with root-detritus than P-acquiring enzymes (acid and alkaline phosphomonoesterase). P fertilisation increased the hotspot areas of C-acquiring enzyme activities over the experiment, without influencing their temporal dynamics. However, its effect on phosphomonoesterase activities depended on the decomposition and delayed the appearance of the highest hotspot areas. P supply met the microbial demand in P-fertilised soil, with high C/P acquisition ratio and constant stoichiometry of microbial biomass C (MBC)/microbial biomass P (MBP). A low C/P acquisition ratio and high MBC/MBP in non-fertilised soil was observed, indicating P limitation for microorganisms. After the 150-day incubation, Olsen P significantly increased in P-fertilised soil (P < 0.05), whereas it decreased in the root-detritusphere of non-fertilised soil. We conclude that the decomposition of root-detritus with a low C/P ratio has potential to improve soil P availability; however, C-P imbalance may increase during the decomposition of root-detritus with a high C/P ratio.

Warming rate drives microbial nutrient demand and enzyme expression during peat decomposition

Recent developments of enzyme-based decomposition models highlight the importance of enzyme kinetics with warming, but most modeling exercises are based on studies with a step-wise warming. This approach may mask the effect of temperature in controlling in-situ activities as in most ecosystems the rate of warming is more gradual than these step warming studies. We conducted an experiment to test the effects of contrasting warming rates on the kinetics of carbon (C), nitrogen (N), and phosphorus (P) degradation enzymes in subtropical peat soils. We also wanted to evaluate if the stoichiometry of enzyme kinetics shifts under contrasting warming rates and if so, how does it relate to the stoichiometry in microbial biomass. Contrasting warming rates altered microbial biomass stoichiometry leading to differing patterns of microbial demand for C vs. nutrient (N and P) and enzyme expression following the optimum foraging strategy. Activity (higher Vmax) and efficiency (lower Km) of C acquisition enzymes were greater in the step treatment; however, expressions of nutrient (N and P) acquiring enzymes were enhanced in the ramp treatment at the end of the experiment. In the step treatment, there was a typical pattern of an initial peak in the Vmax and drop in the Km for all enzyme groups followed by later adjustments. On the other hand, a consistent increase in Vmax and decline in Km of all enzyme groups were observed in the ramp treatment. These changes were sufficient to alter microbial identity (as indicated by enzyme Km and biomass stoichiometry) with two apparently different endpoints under contrasting warming rates. This observation resembles the concept of alternate stable states and highlights a need for improved representation of warming effects on enzymes in decomposition models. Using peat soils of Florida Everglades, here we have demonstrated that contrasting warming rates can influence the dynamics of microbial and enzymatic kinetics. Hence, we suggest that future laboratory and field warming studies could consider our approach to accurately represent microbial and enzymatic kinetics in biogeochemical models.

Responses mechanism of C:N:P stoichiometry of soil microbial biomass and soil enzymes to climate change.

Microorganisms and soil enzymes are important drivers for biogeochemical cycles in terrestrial ecosystems. Understanding the role of microorganisms in the regulation of ecosystems and the response mechanisms of microbial biomass and soil enzymes to climate change are important topic in ecology. From the perspective of climatic factors, this review introduced the roles of microorganisms and soil enzymes in the carbon, nitrogen and phosphorus cycles of terrestrial ecosystems based on the theory of ecological stoichiometry. Moreover, we synthesized the responses mechanisms of soil microbial and soil enzyme stoichiometry, i.e., changes of microbial metabolic rate, enzymatic acti-vity, microbial community structure, ecological stoichiometry of soil microbial biomass and soil enzymes, and nutrient use efficiency. Finally, we analyzed the current research inadequacies and proposed the scientific problems in this field, i.e., to comprehensively elucidate the response mecha-nism of soil microbes and soil enzymes to climate change; to examine the nutrient coupling mechanism of soil microbes and extracellular enzymes; and to explore the adaptive strategies of C:N:P stoichiometry of soil microbial biomass and soil enzymes to climate change.

Microbial stoichiometric flexibility regulates rice straw mineralization and its priming effect in paddy soil

Nitrogen (N) and phosphorus (P) availability plays a crucial role in carbon (C) cycling in terrestrial ecosystems. However, the C:N:P stoichiometric regulation of microbial mineralization of plant residues and its impact on the soil priming effect (PE), measured as CO2 and CH4 emission, in paddy soils remain unclear. In this study, the effect of soil C:N:P stoichiometry (regulated by the application of N and P fertilizers) on the mineralization of 13C-labelled rice straw and the subsequent PE was investigated in a 100-day incubation experiment in flooded paddy soil. N and P additions increased straw mineralization by approximately 25% and 10%, respectively. Additions of both N and P led to higher CO2 efflux, but lower CH4 emission. With an increase in the ratios of DOC:NH4+-N, DOC:Olsen P, and microbial biomass C:N, 13CO2 efflux increased exponentially to a maximum. Compared with sole straw addition, exclusive N addition led to a weaker PE for CO2 emission, whereas exclusive P addition induced a stronger PE for CO2 emission. In contrast, CH4 emitted from native soil organic matter (SOM) was reduced by 7.4% and 46.1% following P and NP application, respectively. Structural equation models suggest that available N had dominant and direct positive effects, whereas microbial biomass stoichiometry mainly exerted negative indirect effects on PE. The stoichiometry of soil enzyme activity directly down-regulated CH4 emission from SOM. Microbes obviously regulate soil C turnover via stoichiometric flexibility to maintain an elemental stoichiometric balance between resources and microbial requirements. The addition of straw in combination with N and P fertilization in paddy soils could therefore meet microbial stoichiometric requirements and regulate microbial activity and extracellular enzyme production, resulting in co-metabolism of fresh C and native SOM.

Effects of vegetation and topography features on ecological stoichiometry of soil and soil microbial biomass in the hilly-gully region of the Loess Plateau, China.

The objectives of this study were to explore the effects of vegetation type, topographic features and their combined effects on soil microbial biomass stoichiometry, so as to better understand the interaction of soil, soil microbes and nutrient cycling under different vegetation types in the hilly-gully region of the Loess Plateau. Soils from three vegetation zones (forest zone, forest-steppe and steppe) and five slope positions (south/north backslope, south/north shoulder and summit) were chosen and the effects of vegetation types and topography features on soil and C:N:P ratios in soil microbial biomass were investigated in this study. The results showed that, among the five slope positions, the highest concentrations of soil and soil microbial biomass C, N, P were found at the backslope position and the north-facing slope. The effects of vegetation types on soil and soil microbial biomass C, N and P in the two soil layers were significantly different, whereas the effects of slope aspect and positions were only numerically different. As for different soil layers, the topsoil (0-10 cm) was more affected by slope aspect, while the subsoil (10-20 cm) was more influenced by slop position. While the effects of vegetation type on soil C:N, C:P and N:P and soil microbial biomass C:N, C:P were significant, slope aspect and slope position only influenced soil C:P and N:P. Consequently, on the Loess Plateau, the effects of vegetation type on soil and soil microbial biomass C, N, P were stronger than those of the topographic features. The standardized major axis tests showed that C:N:P stoichiometry in soil microbes was well-constrained, especially in the steppe zone. The soil microbial biomass N:P might be used as a useful tool to assess nutrient limitation of ecosystem processes in terrestrial ecosystems. If combined with plant leaf N:P, they could provide more accurate information to estimate the nutrient limitation of fragile ecosystem in hilly-gully region of the Loess Plateau.

Chronic Nitrogen Enrichment at the Watershed Scale Does Not Enhance Microbial Phosphorus Limitation

Increased N inputs through chronic atmospheric deposition has enriched temperate forest ecosystems, altering critical ecosystem functions such as decomposition and potentially resulting in a shift to P limitation. We used a combination of microbial biomass stoichiometry and enzymatic activity analyses to evaluate the potential for microbial nutrient limitation over the course of a growing season in response to multi-decadal, whole-watershed N enrichments and a one time, plot-scale P addition that occurred in the 22nd year of whole-watershed treatments. The one-time P addition increased microbial biomass threefold and reduced N-acetyl-glucosaminidase (NAG) and acid phosphatase (AP) activity 1 week after application, but there was no interaction with long-term experimental N enrichment to indicate a shift to P limitation. However, both N and P treatments increased C limitation independently of each other over the duration of the study based on measured increases in β-1,4-glucosidase (BG) activity relative to NAG and AP. Microbial biomass stoichiometry and enzyme activity indicated that BBWM is P limited regardless of N status. Our findings highlight the complex interactions between C, N, and P use and limitation in a forested ecosystem subjected to long-term N enrichment.

Effects of three global change drivers on terrestrial C:N:P stoichiometry: a global synthesis.

Over the last few decades, there has been an increasing number of controlled-manipulative experiments to investigate how plants and soils might respond to global change. These experiments typically examined the effects of each of three global change drivers [i.e., nitrogen (N) deposition, warming, and elevated CO2 ] on primary productivity and on the biogeochemistry of carbon (C), N, and phosphorus (P) across different terrestrial ecosystems. Here, we capitalize on this large amount of information by performing a comprehensive meta-analysis (>2000 case studies worldwide) to address how C:N:P stoichiometry of plants, soils, and soil microbial biomass might respond to individual vs. combined effects of the three global change drivers. Our results show that (i) individual effects of N addition and elevated CO2 on C:N:P stoichiometry are stronger than warming, (ii) combined effects of pairs of global change drivers (e.g., N addition+elevated CO2 , warming+elevated CO2 ) on C:N:P stoichiometry were generally weaker than the individual effects of each of these drivers, (iii) additive interactions (i.e., when combined effects are equal to or not significantly different from the sum of individual effects) were more common than synergistic or antagonistic interactions, (iv) C:N:P stoichiometry of soil and soil microbial biomass shows high homeostasis under global change manipulations, and (v) C:N:P responses to global change are strongly affected by ecosystem type, local climate, and experimental conditions. Our study is one of the first to compare individual vs. combined effects of the three global change drivers on terrestrial C:N:P ratios using a large set of data. To further improve our understanding of how ecosystems might respond to future global change, long-term ecosystem-scale studies testing multifactor effects on plants and soils are urgently required across different world regions.

Soil microbial biomass and function are altered by 12 years of crop rotation

Abstract. Declines in plant diversity will likely reduce soil microbial biomass, alter microbial functions, and threaten the provisioning of soil ecosystem services. We examined whether increasing temporal plant biodiversity in agroecosystems (by rotating crops) can partially reverse these trends and enhance soil microbial biomass and function. We quantified seasonal patterns in soil microbial biomass, respiration rates, extracellular enzyme activity, and catabolic potential three times over one growing season in a 12-year crop rotation study at the W. K. Kellogg Biological Station LTER. Rotation treatments varied from one to five crops in a 3-year rotation cycle, but all soils were sampled under a corn year. We hypothesized that crop diversity would increase microbial biomass, activity, and catabolic evenness (a measure of functional diversity). Inorganic N, the stoichiometry of microbial biomass and dissolved organic C and N varied seasonally, likely reflecting fluctuations in soil resources during the growing season. Soils from biodiverse cropping systems increased microbial biomass C by 28–112 % and N by 18–58 % compared to low-diversity systems. Rotations increased potential C mineralization by as much as 53 %, and potential N mineralization by 72 %, and both were related to substantially higher hydrolase and lower oxidase enzyme activities. The catabolic potential of the soil microbial community showed no, or slightly lower, catabolic evenness in more diverse rotations. However, the catabolic potential indicated that soil microbial communities were functionally distinct, and microbes from monoculture corn preferentially used simple substrates like carboxylic acids, relative to more diverse cropping systems. By isolating plant biodiversity from differences in fertilization and tillage, our study illustrates that crop biodiversity has overarching effects on soil microbial biomass and function that last throughout the growing season. In simplified agricultural systems, relatively small increases in crop diversity can have large impacts on microbial community size and function, with cover crops appearing to facilitate the largest increases.

Stoichiometry of microbial carbon use efficiency in soils

AbstractThe carbon use efficiency (CUE) of microbial communities partitions the flow of C from primary producers to the atmosphere, decomposer food webs, and soil C stores. CUE, usually defined as the ratio of growth to assimilation, is a critical parameter in ecosystem models, but is seldom measured directly in soils because of the methodological difficulty of measuring in situ rates of microbial growth and respiration. Alternatively, CUE can be estimated indirectly from the elemental stoichiometry of organic matter and microbial biomass, and the ratios of C to nutrient‐acquiring ecoenzymatic activities. We used this approach to estimate and compare microbial CUE in &gt;2000 soils from a broad range of ecosystems. Mean CUE based on C:N stoichiometry was 0.269 ± 0.110 (mean ± SD). A parallel calculation based on C:P stoichiometry yielded a mean CUE estimate of 0.252 ± 0.125. The mean values and frequency distributions were similar to those from aquatic ecosystems, also calculated from stoichiometric models, and to those calculated from direct measurements of bacterial and fungal growth and respiration. CUE was directly related to microbial biomass C with a scaling exponent of 0.304 (95% CI 0.237–0.371) and inversely related to microbial biomass P with a scaling exponent of −0.234 (95% CI −0.289 to −0.179). Relative to CUE, biomass specific turnover time increased with a scaling exponent of 0.509 (95% CI 0.467–0.551). CUE increased weakly with mean annual temperature. CUE declined with increasing soil pH reaching a minimum at pH 7.0, then increased again as soil pH approached 9.0, a pattern consistent with pH trends in the ratio of fungal : bacteria abundance and growth. Structural equation models that related geographic variables to CUE component variables showed the strongest connections for paths linking latitude and pH to β‐glucosidase activity and soil C:N:P ratios. The integration of stoichiometric and metabolic models provides a quantitative description of the functional organization of soil microbial communities that can improve the representation of CUE in microbial process and ecosystem simulation models.

Stoichiometry of soil extracellular enzyme activity along a climatic transect in temperate grasslands of northern China

Ratios of specific carbon (C), nitrogen (N) and phosphorus (P) acquisition activities converged on 1:1:1 at a global scale and in tropical ecosystems. It is less clear if this pattern can be applied in temperate grasslands. The questions of whether the pattern remains stable across different soil depths and what the relative contributions are from the influence of climatic, edaphic abiotic and biotic factors on soil extracellular enzyme activity (EEA) stoichiometry remain uncertain. We measured potential activities of one C-acquiring enzyme (β-1, 4-glucosidase), two N-acquiring enzymes (β-N-acetylglucosaminidase and leucine aminopeptidase) and one organic P-acquiring enzyme (acid phosphatase) and major influential factors along a climatic transect in temperate grasslands of northern China during the growing season of 2013. We found lower enzyme C: N (0.47) and C: P (0.18) ratios and a higher enzyme N: P activity ratio (0.40) in 0–20 cm soil depth in temperate grasslands than in tropical soils (C: N, 1.83; C:P, 0.21; N: P, 0.13). The enzyme C: N and C: P ratios decreased with soil depth except for the enzyme C: N ratio in desert steppes. However, there were no significant differences in enzyme N: P ratio with soil depth. Among all the factors, soil total C, N and P contents accounted for the most variation in soil EEA and ecoenzymatic stoichiometry in 0–10 cm surface soil, which implied that soil EEA stoichiometry was largely controlled by soil nutrient stoichiometry. Moreover, edaphic factors had less influence on soil EEA stoichiometry in subsoil than in surface soil and edaphic abiotic factors had a larger effect on soil EEA stoichiometry than climatic and biotic factors. Our results suggest that soil extracellular enzyme activity ratios were not in homeostasis but resource dependent on soil and microbial biomass stoichiometry.

Thermodynamic constraints on the utility of ecological stoichiometry for explaining global biogeochemical patterns

Carbon and nitrogen cycles are coupled through both stoichiometric requirements for microbial biomass and dissimilatory metabolic processes in which microbes catalyse reduction-oxidation reactions. Here, we integrate stoichiometric theory and thermodynamic principles to explain the commonly observed trade-off between high nitrate and high organic carbon concentrations, and the even stronger trade-off between high nitrate and high ammonium concentrations, across a wide range of aquatic ecosystems. Our results suggest these relationships are the emergent properties of both microbial biomass stoichiometry and the availability of terminal electron acceptors. Because elements with multiple oxidation states (i.e. nitrogen, manganese, iron and sulphur) serve as both nutrients and sources of chemical energy in reduced environments, both assimilative demand and dissimilatory uses determine their concentrations across broad spatial gradients. Conceptual and quantitative models that integrate rather than independently examine thermodynamic, stoichiometric and evolutionary controls on biogeochemical cycling are essential for understanding local to global biogeochemical patterns.

Do microorganism stoichiometric alterations affect carbon sequestration in paddy soil subjected to phosphorus input?

Ecological stoichiometry provides a powerful tool for integrating microbial biomass stoichiometry with ecosystem processes, opening far-reaching possibilities for linking microbial dynamics to soil carbon (C) metabolism in response to agricultural nutrient management. Despite its importance to crop yield, the role of phosphorus (P) with respect to ecological stoichiometry and soil C sequestration in paddy fields remains poorly understood, which limits our ability to predict nutrient-related soil C cycling. Here, we collected soil samples from a paddy field experiment after seven years of superphosphate application along a gradient of 0, 30, 60, and 90 (P-0 through P-90, respectively) kg.ha-1.yr-1 in order to evaluate the role of exogenous P on soil C sequestration through regulating microbial stoichiometry. P fertilization increased soil total organic C and labile organic C by 1-14% and 4-96%, respectively, while rice yield is a function of the activities of soil β-1,4-glucosidase (BG), acid phosphatase (AP), and the level of available soil P through a stepwise linear regression model. P input induced C limitation, as reflected by decreases in the ratios of C:P in soil and microbial biomass. An eco-enzymatic ratio indicating microbial investment in C vs. P acquisition, i.e., ln(BG): ln(AP), changed the ecological function of microbial C acquisition, and was stoichiometrically related to P input. This mechanism drove a shift in soil resource availability by increasing bacterial community richness and diversity, and stimulated soil C sequestration in the paddy field by enhancing C-degradation-related bacteria for the breakdown of plant-derived carbon sources. Therefore, the decline in the C:P stoichiometric ratio of soil microorganism biomass under P input was beneficial for soil C sequestration, which offered a "win-win" relationship for the maximum balance point between C sequestration and P availability for rice production in the face of climate change.

Soil microbial biomass C:N:P stoichiometry and microbial use of organic phosphorus

Microbial mineralization and immobilization of nutrients strongly influence soil fertility. We studied microbial biomass stoichiometry, microbial community composition, and microbial use of carbon (C) and phosphorus (P) derived from glucose-6-phosphate in the A and B horizons of two temperate Cambisols with contrasting P availability. In a first incubation experiment, C, nitrogen (N) and P were added to the soils in a full factorial design. Microbial biomass C, N and P concentrations were analyzed by the fumigation-extraction method and microbial community composition was analyzed by a community fingerprinting method (automated ribosomal intergenic spacer analysis, ARISA). In a second experiment, we compared microbial use of C and P from glucose-6-phosphate by adding 14C or 33P labeled glucose-6-phosphate to soil. In the first incubation experiment, the microbial biomass increased up to 30-fold due to addition of C, indicating that microbial growth was mainly C limited. Microbial biomass C:N:P stoichiometry changed more strongly due to element addition in the P-poor soils, than in the P-rich soils. The microbial community composition analysis showed that element additions led to stronger changes in the microbial community in the P-poor than in the P-rich soils. Therefore, the changed microbial biomass stoichiometry in the P-poor soils was likely caused by a shift in the microbial community composition. The total recovery of 14C derived from glucose-6-phosphate in the soil microbial biomass and in the respired CO2 ranged between 28.2 and 37.1% 66 h after addition of the tracer, while the recovery of 33P in the soil microbial biomass was 1.4–6.1%. This indicates that even in the P-poor soils microorganisms mineralized organic P and took up more C than P from the organic compound. Thus, microbial mineralization of organic P was driven by microbial need for C rather than for P. In conclusion, our experiments showed that (i) the microbial biomass stoichiometry in the P-poor soils was more susceptible to additions of C, N and P than in the P-rich soils and that (ii) even in the P-poor soils, microorganisms were C-limited and the mineralization of organic P was mainly driven by microbial C demand.

Linking soil microbial communities to vascular plant abundance along a climate gradient.

The ongoing expansion of shrub cover in response to climate change represents a unique opportunity to explore the link between soil microbial communities and vegetation changes. This link is particularly important in peatlands where shrub expansion is expected to feed back negatively on the carbon sink capacity of these ecosystems. Microbial community structure and function were measured seasonally in four peatlands located along an altitude gradient representing a natural gradient of climate and associated vascular plant abundance. We show that increased soil temperature and reduced water content are associated with greater vascular plant biomass, in particular that of ericoids, and that this, in turn, is correlated with greater microbial biomass. More specifically, microbial community structure is characterized by an increasing dominance of fungi over bacteria with improved soil oxygenation. We also found that the carbon and nitrogen stoichiometry of microbial biomass differs in relation to soil microbial community structure and that this is ultimately associated with a different investment in extracellular enzymatic activity. Our findings highlight the fact that the determination of the structural identity of microbial communities can help to explain the biogeochemical dynamics of organic matter and provide a better understanding of ecosystem response to environmental changes.

Microbial and microfaunal communities in phosphorus limited, grazed grassland change composition but maintain homeostatic nutrient stoichiometry

Previous results from a long-term grassland trial, located in south-east Ireland indicated conserved (homeostatic) nutrient stoichiometry of the soil microbial biomass despite widely varying soil C:N:P ratios. To determine whether this was associated with a change in microbial community structure, rather than a change in microbial physiology, this study characterized the responses of below-ground microbial and nematode community structure to P fertilization. The trial site maintained a range of P fertilisation rates (0–30 kg P ha−1 yr−1) which had been applied since 1968 and soil samples were collected in spring 2009, autumn 2010 and spring 2011. The microbial biomass demonstrated homeostatic stoichiometry over all sampling occasions, particularly of the C:P ratio, despite a 50-fold difference in soil solution C:P ratio. However, microbial and nematode community structure also varied with P fertilisation, indicating that nutrient ratios are maintained even though there were changes in microbial community structure. P fertilization induced a shift from fungal to bacterial dominated decomposition pathways, as indicated by the proportion of bacterial-feeding to fungal-feeding nematodes and bacterial: fungal phospholipid fatty acids (PLFAs). The altered microbial community structure was considered to result from bottom-up control of nutrient quality and quantity by altered vegetation structure and fertilizer inputs, as well as top-down pressures from the nematode community.

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.

Ecoenzymatic Stoichiometry and Ecological Theory

The net primary production of the biosphere is consumed largely by microorganisms, whose metabolism creates the trophic base for detrital foodwebs, drives element cycles, and mediates atmospheric composition. Biogeochemical constraints on microbial catabolism, relative to primary production, create reserves of detrital organic carbon in soils and sediments that exceed the carbon content of the atmosphere and biomass. The production of organic matter is an intracellular process that generates thousands of compounds from a small number of precursors drawn from intermediary metabolism. Osmotrophs generate growth substrates from the products of biosynthesis and diagenesis by enzyme-catalyzed reactions that occur largely outside cells. These enzymes, which we define as ecoenzymes, enter the environment by secretion and lysis. Enzyme expression is regulated by environmental signals, but once released from the cell, ecoenzymatic activity is determined by environmental interactions, represented as a kinetic cascade, that lead to multiphasic kinetics and large spatiotemporal variation. At the ecosystem level, these interactions can be viewed as an energy landscape that directs the availability and flow of resources. Ecoenzymatic activity and microbial metabolism are integrated on the basis of resource demand relative to environmental availability. Macroecological studies show that the most widely measured ecoenzymatic activities have a similar stoichiometry for all microbial communities. Ecoenzymatic stoichiometry connects the elemental stoichiometry of microbial biomass and detrital organic matter to microbial nutrient assimilation and growth. We present a model that combines the kinetics of enzyme activity and community growth under conditions of multiple resource limitation with elements of metabolic and ecological stoichiometry theory. This biogeochemical equilibrium model provides a framework for comparative studies of microbial community metabolism, the principal driver of biogeochemical cycles.

Production, partitioning and stoichiometry of organic matter under variable nutrient supply during mesocosm experiments in the tropical Pacific and Atlantic Ocean

Abstract. Oxygen-deficient waters in the ocean, generally referred to as oxygen minimum zones (OMZ), are expected to expand as a consequence of global climate change. Poor oxygenation is promoting microbial loss of inorganic nitrogen (N) and increasing release of sediment-bound phosphate (P) into the water column. These intermediate water masses, nutrient-loaded but with an N deficit relative to the canonical N:P Redfield ratio of 16:1, are transported via coastal upwelling into the euphotic zone. To test the impact of nutrient supply and nutrient stoichiometry on production, partitioning and elemental composition of dissolved (DOC, DON, DOP) and particulate (POC, PON, POP) organic matter, three nutrient enrichment experiments were conducted with natural microbial communities in shipboard mesocosms, during research cruises in the tropical waters of the southeast Pacific and the northeast Atlantic. Maximum accumulation of POC and PON was observed under high N supply conditions, indicating that primary production was controlled by N availability. The stoichiometry of microbial biomass was unaffected by nutrient N:P supply during exponential growth under nutrient saturation, while it was highly variable under conditions of nutrient limitation and closely correlated to the N:P supply ratio, although PON:POP of accumulated biomass generally exceeded the supply ratio. Microbial N:P composition was constrained by a general lower limit of 5:1. Channelling of assimilated P into DOP appears to be the mechanism responsible for the consistent offset of cellular stoichiometry relative to inorganic nutrient supply and nutrient drawdown, as DOP build-up was observed to intensify under decreasing N:P supply. Low nutrient N:P conditions in coastal upwelling areas overlying O2-deficient waters seem to represent a net source for DOP, which may stimulate growth of diazotrophic phytoplankton. These results demonstrate that microbial nutrient assimilation and partitioning of organic matter between the particulate and the dissolved phase are controlled by the N:P ratio of upwelled nutrients, implying substantial consequences for nutrient cycling and organic matter pools in the course of decreasing nutrient N:P stoichiometry.

C:N:P stoichiometry and nutrient limitation of the soil microbial biomass in a grazed grassland site under experimental P limitation or excess

IntroductionThe availability of essential nutrients, such as nitrogen (N) and phosphorus (P), can feedback on soil carbon (C) and the soil microbial biomass. Natural cycles can be supplemented by agricultural fertiliser addition, and we determined whether the stoichiometry and nutrient limitation of the microbial biomass could be affected by an unbalanced nutrient supply.MethodsSamples were taken from a long-term trial (in effect since 1968) with annual applications of 0, 15 and 30 kg P ha−1 with constant N and potassium. Soil and microbial biomass CNP contents were measured and nutrient limitation assessed by substrate-induced respiration. Linear regression and discriminant analyses were used to identify the variables explaining nutrient limitation.ResultsSoil and biomass CNP increased with increasing P fertiliser, and there was a significant, positive, correlation between microbial biomass P and biomass C, apart from at the highest level of P fertilisation when the microbial biomass was over-saturated with P. The molar ratios of C:N:P in the microbial biomass remained constant (homeostatic) despite large changes in the soil nutrient ratios. Microbial growth was generally limited by C and N, except in soil with no added P when C and P were the main limiting nutrients. C, N and P, however, did not explain all the growth limitation on the soils with no added P.ConclusionsIncreased soil C and N were probably due to increased net primary production. Our results confirm that C:N:P ratios within the microbial biomass were constrained (i.e. homeostatic) under near optimum soil conditions. Soils with no added P were characterised by strong microbial P limitation and soils under high P by over-saturation of microorganisms with P. Relative changes in biomass C:P can be indicative of nutrient limitation within a site.

Effects of Resource Chemistry on the Composition and Function of Stream Hyporheic Biofilms

Fluvial ecosystems process large quantities of dissolved organic matter as it moves from the headwater streams to the sea. In particular, hyporheic sediments are centers of high biogeochemical reactivity due to their elevated residence time and high microbial biomass and activity. However, the interaction between organic matter and microbial dynamics in the hyporheic zone remains poorly understood. We evaluated how variance in resource chemistry affected the microbial community and its associated activity in experimentally grown hyporheic biofilms. To do this we fed beech leaf leachates that differed in chemical composition to a series of bioreactors filled with sediment from a sub-alpine stream. Differences in resource chemistry resulted in differences in diversity and phylogenetic origin of microbial proteins, enzyme activity, and microbial biomass stoichiometry. Specifically, increased lignin, phenolics, and manganese in a single leachate resulted in increased phenoloxidase and peroxidase activity, elevated microbial biomass carbon:nitrogen ratio, and a greater proportion of proteins of Betaproteobacteria origin. We used this model system to attempt to link microbial form (community composition and metaproteome) with function (enzyme activity) in order to better understand the mechanisms that link resource heterogeneity to ecosystem function in stream ecosystems.