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While understanding feeding preferences of herbivores and carnivores is of major importance in ecology, we still know very little on the sensitivity of different functional groups to suboptimal stoichiometric resource quality. Here, we apply concepts of ecological stoichiometry to shed light on differences in the nutritional requirements of herbivores and carnivores, and to make predictions on the influence of suboptimal resource stoichiometric quality on the fitness of these different consumers to. Herbivores generally experience more variation in the quality of their resource than carnivores do, and these differences have likely shaped the extent to which coping mechanisms have evolved. Consequently, we expect 1) herbivores to maintain their stoichiometric homeostasis over a broader range of resource stoichiometry than carnivores, 2) the threshold elemental ratio (TER), i.e. the dietary carbon to nutrient ratio which maximizes fitness, of herbivores to be higher than that of carnivores, 3) a narrower and sharper knife‐edge response in carnivores than herbivores and 4) asymmetric knife‐edge responses indicating a higher sensitivity to the diet quality that consumers are not used to dealing with, namely nutrient limitation in carnivores and nutrient excess in herbivores. Our study poses that documenting the ranges of resource quality where consumer fitness declines in diverse organisms is a very promising avenue to increase our understanding of community composition and food web functioning. [ABSTRACT FROM AUTHOR]

In light of ongoing climate change, it is increasingly important to know how nutritional requirements of ectotherms are affected by changing temperatures. Here, we analyse the wide thermal response of phosphorus (P) requirements via elemental gross growth efficiencies of Carbon (C) and P, and the Threshold Elemental Ratios in different aquatic invertebrate ectotherms: the freshwater model species Daphnia magna, the marine copepod Acartia tonsa, the marine heterotrophic dinoflagellate Oxyrrhis marina, and larvae of two populations of the marine crab Carcinus maenas. We show that they all share a non‐linear cubic thermal response of nutrient requirements. Phosphorus requirements decrease from low to intermediate temperatures, increase at higher temperatures and decrease again when temperature is excessive. This common thermal response of nutrient requirements is of great importance if we aim to understand or even predict how ectotherm communities will react to global warming and nutrient‐driven eutrophication. [ABSTRACT FROM AUTHOR]

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Abstract Background Trait based functional and community ecology is en vogue. Most studies, however, ignore phenotypical diversity by characterizing entire species considering only trait means rather than their variability. Phenotypical variability may arise from genotypical differences or from ecological factors (e.g., nutritionally imbalanced diet), and these causes can usually not be separated in natural populations. We used a single genotype from a parthenogenetic model system (the oribatid mite Archegozetes longisetosus Aoki) to exclude genotypical differences. We investigated patterns of dietary (10 different food treatments) induced trait variation by measuring the response of nine different traits (relating to life history, morphology or exocrine gland chemistry). Results Nutritional quality (approximated by carbon-to-nitrogen ratios) influenced all trait means and their variation. Some traits were more prone to variation than others. Furthermore, the “threshold elemental ratio”- rule of element stoichiometry applied to phenotypic trait variation. Imbalanced food (i.e. food not able to fully meet the nutritional demands of an animal) led to lower trait mean values, but also to a higher variation of traits. Conclusion Imbalanced food led not only to lower trait value averages, but also to higher trait variability. There was a negative relationship between both parameters, indicating a direct link of both, average trait levels and trait variation to nutritional quality. Hence, variation of trait means may be a predictor for general food quality, and further indicate trade-offs in specific traits an animal must deal with while feeding on imbalanced diets.

1. All bark beetles are in symbiosis with fungi. Although obligate mutualisms with fungi are common with tree‐killing bark beetles (primaries), fungi associated with non‐tree‐killing bark beetles (secondaries) are usually dismissed as commensals. 2. Using an ecological stoichiometric approach, we show secondaries are also involved in nutrition‐based mutualisms, some of which appear obligate, and that differences in symbiont provisioning efficiency have a potent effect on beetle carbon (C): nitrogen (N): phosphorus (P) ratios. 3. Some secondary beetles have high P contents and require efficient P provisioning via fungi, while others have low P contents that may allow them to exploit less efficient fungi or a broader range of species with variable efficiencies. A co‐occurring scavenger that feeds on nutrient‐poor bark beetle frass (excrement/boring residues) exhibited the lowest phosphorus content yet recorded for an invertebrate. 4. Our results generally support the growth‐rate hypothesis that posits differences in C:P and N:P ratios in consumers are due to differential allocation of P to P‐rich RNA to support growth. However, while the beetle species that accumulated the most biomass was considerably enriched in P and that with the least biomass was P‐poor, one beetle species that was P‐rich was also small possibly due to limitation by an element other than P. 5. Our results indicate that fungi are important to a broader range of bark beetles than previously recognised. Additional research is needed to describe how these various symbioses influence forest ecosystems via differential effects of fungi on host beetle fitness. [ABSTRACT FROM AUTHOR]

1. Ecological stoichiometry theory was applied to investigate how a consumer contends with an extreme elemental mismatch between its food and its body via symbiotic facilitation. 2. The beetle Dendroctonus brevicomis LeConte develops in bark, a substrate extremely low in nitrogen (N) and phosphorus (P). Its survival there depends on interactions with mutualist and antagonist fungi. 3. This study found that mutualists transfer N and P from sapwood and phloem into bark, where beetles feed, whereas the antagonist moves these elements only to phloem, resulting in starvation of the insect. However, even with mutualists, N and P concentrations remained low in bark, resulting in low N and extremely low P concentrations in the beetle. 4. The N:P ratios found in D. brevicomis larvae were the highest thus far reported for beetles and among the highest for insects and invertebrates. This suggests that the beetle has evolved additional, nutrient‐sparing adaptations. [ABSTRACT FROM AUTHOR]

Abstract Global nitrogen (N) deposition can change the contents and stoichiometry of soil resources and thus of microbial communities, which can drive the flow of carbon (C) and nutrients in food webs in forest ecosystems. It is critical to understand the status of soil elements required for microbial growth, the elements of microbial growth, and how soil microorganisms would reallocate resources and release extracellular enzymes for adjusting the imbalance between resources and microorganisms due to N deposition. We chose a plantation of Pinus tabuliformis where N had been added across a gradient (0–9 g N m−2 y−1) for two years to reveal how matters flow and soil resources were reallocated by microbes to cope N addition. Our results showed that although two years of N inputs only significantly affect the ratio between the activities of β-1,4- N ‑acetylglucosaminidase and alkaline phosphatase in the 0–20 cm soil layer, activities for both organic N and organic phosphorus (P) acquisition enzymes scale with C acquisition with slope of about 1, following the global ecoenzymatic patterns. Contents and elemental ratios of soil organic carbon (SOC), total N (TN) and total phosphorus (TP) were changed after this short-term N addition. Ratios of SMBC, SMBN and SMBP were concentrated from soil and lower than ratios of soil resources. Soil microbial communities under N addition were limited by N and co-limited by P at 3 and 6 g N m−2 y−1. Although imbalance between soil and microbes caused by N addition, in the 0–20 and 20–40 cm layers, C: N relationship between soil and the microbial community indicated microbial community maintained homeostasis. However, C:P relationship between soil and the microbial community indicated no microbial community homeostasis. In addition, both soil resources elemental stoichiometry and contents can affect soil microbial communities. Further study of the structure of soil microbial communities, identifying the microbes that are more adaptive to high N concentrations and N deposition, is needed for a better understanding of the mechanisms of nutrient flow in the food web and how to maintain microbial homeostasis in response to N deposition in forest ecosystems. Highlights • Enzymatic activities, microbial coping with N addition, maintain no change. • Microbial communities are limited by N and co-limited by P at N3 and N6. • Soil microbes generally maintain community-level elemental homeostasis. • Contents and stoichiometry of soil resources influence microbial communities. [ABSTRACT FROM AUTHOR]

Stoichiometric ratios of resources and consumers have been used to predict nutrient limitation across diverse terrestrial and aquatic ecosystems. In forested headwater streams, coarse and fine benthic organic matter (CBOM, FBOM) are primary basal resources for the food web, and the distribution and quality of these organic matter resources may therefore influence patterns of secondary production and nutrient cycling within stream networks or among biomes. We measured carbon (C), nitrogen (N), and phosphorus (P) content of CBOM and FBOM and calculated their stoichiometric ratios (C/N, C/P, N/P) from first- to fourth-order streams from tropical montane, temperate deciduous, and boreal forests, and tallgrass prairie, to compare the magnitude and variability of these resource types among biomes. We then used the ratios to predict nutritional limitations for consumers of each resource type. Across biomes, CBOM had consistently higher %C and %N, and higher and more variable C/N and C/P than FBOM, suggesting that microbial processing results in more tightly constrained elemental composition in FBOM than in CBOM. Biome-specific differences were observed in %P and N/P between the two resource pools; CBOM was lower in %P but higher in N/P than FBOM in the tropical montane and temperate deciduous forest biomes, while CBOM was higher in %P but similar in N/P than FBOM in the grassland and boreal forest biomes. Stable 13C isotopes suggest that FBOM likely derives from CBOM in tropical and temperate deciduous forest, but that additional non-detrital components may contribute to FBOM in boreal forests and grasslands. Comparisons of stoichiometric ratios of CBOM and FBOM to estimated needs of aquatic detritivores suggest that shredders feeding on CBOM are more likely to experience nutrient (N and/or P) than C limitation, whereas collector-gatherers consuming FBOM are more likely to experience C than N and/or P limitation. Our results suggest that differences in basal resource elemental content and stoichiometric ratios have the potential to affect consumer production and ecosystem rates of C, N, and P cycling in relatively consistent ways across diverse biomes. [ABSTRACT FROM AUTHOR]

Abstract: Quantitative approaches to predator–prey interactions are central to understanding the structure of food webs and their dynamics. Different predatory strategies may influence the occurrence and strength of trophic interactions likely affecting the rates and magnitudes of energy and nutrient transfer between trophic levels and stoichiometry of predator–prey interactions. Here, we used spider–prey interactions as a model system to investigate whether different spider web architectures—orb, tangle, and sheet‐tangle—affect the composition and diet breadth of spiders and whether these, in turn, influence stoichiometric relationships between spiders and their prey. Our results showed that web architecture partially affects the richness and composition of the prey captured by spiders. Tangle‐web spiders were specialists, capturing a restricted subset of the prey community (primarily Diptera), whereas orb and sheet‐tangle web spiders were generalists, capturing a broader range of prey types. We also observed elemental imbalances between spiders and their prey. In general, spiders had higher requirements for both nitrogen (N) and phosphorus (P) than those provided by their prey even after accounting for prey biomass. Larger P imbalances for tangle‐web spiders than for orb and sheet‐tangle web spiders suggest that trophic specialization may impose strong elemental constraints for these predators unless they display behavioral or physiological mechanisms to cope with nutrient limitation. Our findings suggest that integrating quantitative analysis of species interactions with elemental stoichiometry can help to better understand the occurrence of stoichiometric imbalances in predator–prey interactions. [ABSTRACT FROM AUTHOR]

Arid ecosystems are characterized as having stressful conditions of low energy and nutrient availability for soil microorganisms and vegetation. The rhizosphere serves as the one of most active microorganism habitats, however, the general understanding of the ecoenzymatic stoichiometry (exoenzymes) and microbial nutrient acquisition in rhizosphere soil is limited. Here, we investigated the vegetation communities and determined the soil physicochemical properties, microbial biomass, and enzymatic activities in rhizosphere under different vegetation and soil types in the arid area of the northern Loess Plateau. Type Ⅱ standard major axis (SMA) regression analysis showed that the plants played a more important role than soil properties in determining ecoenzymatic stoichiometry. Linear regression analysis displayed a microbial stoichiometric homeostasis (community-level) in rhizosphere. The Threshold Elemental Ratio (TER) revealed that the microbial nutrient metabolisms of rhizosphere were co-limited by N and P in the A. ordosica and A. cristatum communities of loess, and A. cristatum communities of feldspathic sandstone weathered soil. Binding spatial ordination analysis (RDA and CCA) demonstrated that soil physical properties (e.g., soil moisture, silt and clay contents) have more contribution to ecoenzymatic stoichiometry than the other investigated soil parameters, whereas soil nutrients (e.g., total organic carbon, nitrogen, and phosphorus) predominantly controlled microbial nutrient ratios. Therefore, the ecoenzymatic stoichiometry in rhizosphere is greatly regulated by plants and soil physical properties. The microbial N and P are co-limited under Gramineae plant in loess and feldspathic sandstone weathered soil regions. Meanwhile, the microbial nutrient limitation is mainly affected by soil nutrient supply. These findings could be crucial for illuminating rhizosphere microbial metabolism and revealing the nutrient cycling of root-soil interface under arid and oligotrophic ecosystems. [ABSTRACT FROM AUTHOR]

Background Soil microbial communities (SMC) play a central role in the structure and function of desert ecosystems. However, the high variability of annual precipitation could results in the alteration of SMC and related biological processes depending on soil water potential. The nature of the physiological adjustments made by SMC in order to obtain energy and nutrients remains unclear under different soil resource availabilities in desert ecosystems. In order to examine this dynamic, the present study examined the effects of variation in annual precipitation on physiological adjustments by the SMC across two vegetation-soil systems of different soil organic matter input in an oligotrophic desert ecosystem. Methods We collected soil samples in the Cuatro Ciénegas Basin (Mexico) under two vegetation covers: rosetophylous scrub (RS) and grassland (G), that differ in terms of quantity and quality of organic matter. Collections were conducted during the years 2011, 2012, 2013 and 2014, over which a noticeable variation in the annual precipitation occurred. The ecoenzymatic activity involved in the decomposition of organic matter, and the concentration of dissolved, available and microbial biomass nutrients, were determined and compared between sites and years. Results In 2011, we observed differences in bacterial taxonomic composition between the two vegetation covers. The lowest values of dissolved, available and microbial nutrients in both cover types were found in 2012. The G soil showed higher values of dissolved and available nutrients in the wet years. Significant positive correlations were detected between precipitation and the ratios Cmic:Nmic and Cmic:Pmic in the RS soil and Cmic:Pmic and Nmic:Pmic in the G soil. The slopes of the regression with Cmic and Nmic were higher in the G soil and lower in the RS soil. Moreover, the SMC under each vegetation cover were co-limited by different nutrients and responded to the sum of water stress and nutrient limitation. Discussion Soil community within both sites (RS and G) may be vulnerable to drought. However, the community of the site with lower resources (RS) is well adapted to acquire P resources by ecoenzyme upregulation during years with adequate precipitation, suggesting that this community is resilient after drought occurs. Under the Global Climate Change scenarios for desert ecosystems that predict reduced annual precipitation and an increased intensity and frequency of torrential rains and drought events, the soil microbial communities of both sites could be vulnerable to drought through C and P co-limitation and reallocation of resources to physiological acclimatization strategies in order to survive.

Elevated temperature causes metabolism and respiration to increase in poikilothermic organisms. We hypothesized that invertebrate consumers will therefore require increasingly carbon-rich diets in a warming environment because the increased energetic demands are primarily met using compounds rich in carbon, that is, carbohydrates and lipids. Here, we test this hypothesis using a new stoichiometric model that has carbon (C) and nitrogen (N) as currencies. Model predictions did not support the hypothesis, indicating instead that the nutritional requirements of invertebrates, at least in terms of food quality expressed as C:N ratio, may change little, if at all, at elevated temperature. Two factors contribute to this conclusion. First, invertebrates facing limitation by nutrient elements such as N have, by default, excess C in their food that can be used to meet the increased demand for energy in a warming environment, without recourse to extra dietary C. Second, increased feeding at elevated temperature compensates for the extra demands of metabolism to the extent that, when metabolism and intake scale equally with temperature (have the same Q10), the relative requirement for dietary C and N remains unaltered. Our analysis demonstrates that future climate-driven increases in the C:N ratios of autotroph biomass will likely exacerbate the stoichiometric mismatch between nutrient-limited invertebrate grazers and their food, with important consequences for C sequestration and nutrient cycling in ecosystems. [ABSTRACT FROM AUTHOR]

Arid ecosystems are characterized by stressful conditions of low energy and nutrient availability for soil microorganisms. It has been observed that the ecoenzymes needed for the transformation of organic compounds into assimilable products show similar scaling relationships in different habitats (logarithmic C:N:P scaling ratios ~1:1:1). In this study in Cuatro Ciénegas Basin (CCB) in the Chihuahuan desert of México, we report among the lowest ecoenzymatic activities yet quantified in soil. Nevertheless, activities for both organic N and organic P acquisition enzymes scale with C acquisition with a slope of ∼1.0, indicating that the soil microbial communities of this ultra-oligotrophic desert ecosystem follow the global ecoenzymatic stoichiometry patterns. CCB soil microbial communities were co-limited by C and either by N or P but this co-limitation played out differently in different parts of the CCB as indicated by microbial ecoenzymatic shift to allocate more resources to acquire and immobilize the scarcer nutrient. By extending ecoenzymatic analyses to these ultra-oligotrophic soils, our findings support the broad utility of the approach in illuminating how microbes acquire limiting resources in arid ecosystems. [ABSTRACT FROM AUTHOR]

Temperature and nutrient availability play key roles in controlling the pathways and rates at which energy and materials move through ecosystems. These factors have also changed dramatically on Earth over the past century as human activities have intensified. Although significant effort has been devoted to understanding the role of temperature and nutrients in isolation, less is known about how these two factors interact to influence ecological processes. Recent advances in ecological stoichiometry and metabolic ecology provide a useful framework for making progress in this area, but conceptual synthesis and review are needed to help catalyze additional research. Here, we examine known and potential interactions between temperature and nutrients from a variety of physiological, community, and ecosystem perspectives. We first review patterns at the level of the individual, focusing on four traits - growth, respiration, body size, and elemental content - that should theoretically govern how temperature and nutrients interact to influence higher levels of biological organization. We next explore the interactive effects of temperature and nutrients on populations, communities, and food webs by synthesizing information related to community size spectra, biomass distributions, and elemental composition. We use metabolic theory to make predictions about how population-level secondary production should respond to interactions between temperature and resource supply, setting up qualitative predictions about the flows of energy and materials through metazoan food webs. Last, we examine how temperature-nutrient interactions influence processes at the whole-ecosystem level, focusing on apparent vs. intrinsic activation energies of ecosystem processes, how to represent temperature-nutrient interactions in ecosystem models, and patterns with respect to nutrient uptake and organic matter decomposition. We conclude that a better understanding of interactions between temperature and nutrients will be critical for developing realistic predictions about ecological responses to multiple, simultaneous drivers of global change, including climate warming and elevated nutrient supply. [ABSTRACT FROM AUTHOR]

The formation and development of biological soil crusts (biocrusts) potentially affect the cycles and stoichiometric characteristics of soil carbon (C), nitrogen (N), and phosphorus (P). However, it is still unclear how soil microbes adapt to such changes. In this study, we examined the effects of moss-dominated biocrusts coverage (0, 1%-20%, 20%-40%, 40%-60%, 60%-80%, and 80%-100%) on soil physicochemical properties, soil microbial biomass, and ectoenzyme activities [β-1, 4-glucosidase (BG), β-1, 4-N-acetyl glucosidase (NAG), acid phosphatase (AP)] in two soil layers (0-5 and 5-10 cm) in the Three Gorges Reservoir area, as well as the covariations of soil-microbe-ectoenzyme C:N:P stoichiometry. The results showed that biocrust development significantly increased soil clay content, water stable aggregates, soil C, N, P contents, and significantly decreased soil bulk density and sand content. Microbial biomass C, N, P and ectoenzyme activities were significantly increased with increasing biocrust coverage. Soil depth did not affect soil physicochemical properties and C:N:P, but significantly affected microbial biomass, ectoenzyme activities, BG:AP and NAG:AP. Soil C, N and P contents were significantly positively correlated with microbial biomass and ectoenzyme activities, negatively correlated with BG:NAG, while positively correlated with NAG:AP, but had no significant correlation with microbial biomass C:N:P. There was no significant correlation between soil-microbe and microbial-ectoenzyme C:N:P. BG:NAG:AP decreased gradually with the increase of C:N:P stoichiometric imbalance between microbe and soil. This study indicated that the microbial metabolism was co-limited by N and P and with stronger P limitation. Microbes could maintain homeostasis by adjusting their own biomass and ectoenzyme C:N:P to adapt to changes in soil ecological stoichiometry driven by biocrust development.

1. Ecological stoichiometry expresses ecological interactions as the balance between multiple elements. It relates the ecological function of organisms to their elemental composition, or their organismal stoichiometry. Organismal stoichiometry is thought to reflect elemental investments in life history and morphology acting in concert with variability in abiotic or environmental conditions, but the relative contribution of these factors to natural variability in organismal stoichiometry is poorly understood. 2. We assessed the relative contribution of stream identity, predation, body size and sex to the organismal stoichiometry of guppies ( Poecilia reticulata) in six streams in Trinidad. In this system, guppy life-history phenotype evolves in response to predation. Guppies adapted to high-predation (HP) pressure grow faster, mature earlier, produce fewer and smaller offspring and eat a higher-quality diet than guppies adapted to low-predation (LP) pressure. This pattern of life-history evolution is repeated in many rivers encompassing a wide range of abiotic conditions. 3. Organismal stoichiometry of guppies was widely variable, spanning up to ∼70% of the range of variability reported across freshwater fish taxa. Streams from where guppies were sampled were the most important predictor of organismal stoichiometry. In many cases, guppy populations from sites within the same stream varied as much as from sites in different streams. 4. Surprisingly, predation regime was a minor predictor of % C, C : P and C : N in female guppies, despite its strong correlation with life-history phenotype and other organismal traits in this species. Body size and sex were not significant predictors of organismal stoichiometry. 5. Guppies from HP sites were more stoichiometrically balanced with their diets than guppies from LP sites. The latter appeared to be more vulnerable to phosphorus limitation than the former, suggesting that dietary specialization associated with guppy life-history phenotype may have stoichiometric consequences that can affect guppy physiology and nutrient recycling. 6. Our findings suggest that local environmental conditions are a stronger predictor of organismal stoichiometry than organismal traits. We recommend that future work should explicitly consider correlations between organismal traits and organismal stoichiometry in the context of environmental heterogeneity. [ABSTRACT FROM AUTHOR]

1. The body tissues of insect herbivores contain higher concentrations of nitrogen and phosphorus than do their host plants, leading to an elemental mismatch that can limit herbivore growth, fecundity and ultimately influence population dynamics. While low nutrient availability can limit herbivore growth and reproduction, nutrient levels that exceed an organism’s nutritional requirements, i.e. an organisms’ threshold elemental ratio, can also decrease performance. 2. We conducted a laboratory experiment to examine the impacts of nitrogen and phosphorus additions on population growth rates of a phloem-feeding insect herbivore. 3. Herbivore per capita population growth rates were highest at intermediate foliar nitrogen concentrations, indicating a performance cost on the highest nitrogen foliage. While there was no direct effect of foliar phosphorus concentration on insect performance, there was a strong and unexpected indirect effect. High soil phosphorus availability increased both foliar nitrogen concentrations and aphid tissue nitrogen, resulting in low population growth rates when both soil nitrogen and phosphorus availabilities were high. 4. In this study, experimental increases in foliar nitrogen levels led to a decrease in herbivore performance suggesting that excessive nutrient levels can limit herbivore population growth rates. [ABSTRACT FROM AUTHOR]

Nutritional imbalances are of great interest in the ecological stoichiometry literature, in which researchers have focused almost exclusively on cases where nutrients are available in low amounts relative to energy (carbon), and animal growth is impaired due to insufficient nutrient intake. Little attention has been given to situations where food elemental content is higher than the level that satisfies animal requirements. However, most animals are strongly homeostatic with respect to the elemental composition of their body; hence they must excrete the excess of elements that are not in short supply. To date, stoichiometric theory has assumed that excretion of superfluous elements does not come with a cost and, thus, that consumption of food with surplus nutrients does not impair performance. Here we challenge this assumption, based on a compilation of several examples involving food phosphorus content that show that the performance of a wide variety of animals decreases when supplied with food containing high concentrations of (potentially) limiting nutrients. We discuss possible mechanisms for this phenomenon, and suggest that animals most vulnerable to effects of high food nutrient content are those that normally feed on low- quality (low-nutrient: C) food, and have a relatively low body nutrient content themselves, such as herbivores and detritivores. [ABSTRACT FROM AUTHOR]

1. The transition from carbon (C) to phosphorus (P) limited growth in Daphnia depends not only on the C : P ratio in seston, i.e. food quality, but also on food quantity. Carbon is commonly believed to be limiting at low food because of the energetic demands of basal metabolism. The critical C : P ratio in seston (otherwise known as the threshold elemental ratio, TER) above which P is limiting would then be high when food is scarce. 2. A new model that differentiates between the C : P requirements for growth and maintenance is presented that includes terms for both C and P in basal metabolism. At low food the calculated TERs for Daphnia of around 230 are only slightly higher than values of 200 or so at high intake. Seston C : P often exceeds 230, particularly in oligotrophic lakes where phytoplankton concentration is low and detritus dominates the diet, indicating the potential for limitation by P. 3. The analysis highlights the importance of P, as well as C, in maintenance metabolism and the overall metabolic budget, such that food quality is of importance even when intake is low. Further measurements of C and P metabolism at low food, in particular basal respiration and excretion rates, are needed in order to improve our understanding of the interacting roles of food quantity and quality in zooplankton nutrition. [ABSTRACT FROM AUTHOR]

• Soil EES indicate microbial nutritional status and resource limitations. • Thinning and seasonal changes affect ecoenzymes and microbial metabolism in the soil. • Differences in soil EEA and EES may be related to the variations in soil properties, especially soil nutrients and microbial biomass. • Soil microbial metabolism was limited by N in Larix principis-rupprechtii plantations. The extracellular ecoenzymatic stoichiometry (EES) of the soil are key indicators of the nutritional status and relative resource limitations of microbes that play important roles in biogeochemical cycling and ecosystem functions. However, our knowledge of the effects of forest thinning and the seasons on the variability of extracellular ecoenzymatic activities (EEA) and EES in the soil is limited. We examined the soils from Larix principis-rupprechtii plantations after nine years of treatments with four different thinning intensities (i.e., control (CK), low thinning (LT), moderate thinning (MT), and high thinning (HT)) in four different seasons. We measured the activities of one carbon (C) acquiring enzyme (β-1,4-glucosidase (BG)), two nitrogen (N) acquiring enzymes (β-1,4-N-acetylglucosaminidase (NAG), leucine aminopeptidase (LAP)), and one phosphorus (P) acquiring enzyme (acid phosphatase (AP)), and examined potential factors (soil environment, nutrients, and microbial biomass) that may influence the activities of these enzymes. Our results showed that thinning significantly (p < 0.05) enhanced the activities of the BG, NAG + LAP, and AP. The EEA were higher in summer and autumn than those in spring and winter. Further, MT and HT stands exhibited lower BG:(NAG + LAP) and higher (NAG + LAP):AP ratios than the CK and LT stands, demonstrating that MT and HT caused a greater N limitation for soil microbes. The higher (NAG + LAP):AP ratio during winter indicated that the microbial N limitation was increased in this season. The variations in the EEA, EES, environment, nutrients, and microbial biomass in the soil among different thinning treatments demonstrated that thinning could markedly promote microbial activities and metabolism and improve nutrient cycling. Based on analyses of the BG:(NAG + LAP) ratio, (NAG + LAP):AP ratio, and the threshold element ratio, we found that the metabolism of soil microbes was limited by N in L. principis-rupprechtii plantations in North China. A redundancy analysis and a variation partitioning analysis revealed that variations in the EEA and EES of soils could attributed to the alterations in soil properties, especially soil nutrient and microbial biomass. Overall, the ecoenzymatic C:N:P stoichiometry was dependent on the resource availability in the soils rather than on homeostatic mechanisms. Our research highlights the importance of the effects of thinning practices and season types on ecoenzymes and microbial metabolism, and provides insights into the microbial resource limitation, nutrient cycling, and ecological processes in plantation forests after thinning. [ABSTRACT FROM AUTHOR]

The highland forests of tropical regions are highly vulnerable to climate change because changes in soil organic quality due to the increased soil water deficit conditions through rising temperatures. Several authors have reported that labile molecules dominate soil organic matter at higher elevations, and it is therefore more vulnerable to the rising temperatures associated with climate change. The objective of the present study was to analyze the effect of interaction between the chemical composition of organic matter derived from the dominant plant species and the metabolism of microbial community along an elevational gradient in a highland forest in Central Mexico. The study compared three vegetation-soil systems that represent three different elevational levels: Alnus -system (3100 m.a.s.l.), Abies -system (3500 m.a.s.l.) and Pinus -system (3700 m.a.s.l.). The SOM produced in the lowest site is more recalcitrant (i.e., higher Alkyl : O - Alkyl ratio) as a result of the lower water availability than in the highest site. The results of Threshold Elemental Ratio C:N (TER C:N) and TER C:P for the organic layer were lower than their C:N and C:P ratios in the organic layer, supporting that the microbial community of the organic layer in the site of lowest elevation must be limited by the carbon source, rather than by N and P. However, these results were not found in the mineral soil, suggesting that the drivers of organic matter decomposition differ between the organic layer and the mineral soil. As a conclusion, our results suggest that the chemical recalcitrance of organic matter (at the lowest site) and temperature (at the highest site) reduce the microbial metabolic activity in the forest floor. Integrated study of plant-derived organic material and the microbial metabolism of the forest floor is therefore required to achieve a full understanding of the vulnerability of tropical mountain ecosystems to climate change. Unlabelled Image • The SOM produced by vegetation in the lowest site is more recalcitrant than in a higher elevation site. • The microbial community of the organic layer in the highest elevation site is limited by N and P. • The drivers of organic matter decomposition differ between the organic layer and the mineral soil. [ABSTRACT FROM AUTHOR]

• Microbial processes during secondary succession were studied on the Loess Plateau. • Microbial communities showed stoichiometric homeostasis during plant secondary succession. • Microbial metabolisms were co-limited by C and P during plant secondary succession. • Microbial activities were strongly affected by soil nutrient status. Soil microbial metabolism is vital for nutrient cycling and ecosystem stability. To quantify microbial metabolism and nutrient limitation during plant secondary succession, we measured soil physicochemical properties, microbial biomass, and four enzyme activities (β-1,4-glucosidase (BG), β-1,4- N -acetylglucosaminidase (NAG), L -leucine aminopeptidase (LAP), and alkaline phosphatase (AP)) and identified the changes of soil ecoenzymatic stoichiometry and microbial nutrient limitation along a secondary succession series in typical arid and semi-arid ecosystems on the Loess Plateau, China. Soil enzyme activities increased in the first 17a succession and then decreased with plant secondary succession. The ln(BG):ln(NAG + LAP) ratio and ln(BG):ln(AP) ratio showed a decreasing trend in the first 22a succession and then increased. Most soil nutrient contents and nutrient stoichiometry were significantly correlated with enzyme activities and enzymatic stoichiometry. Moreover, vector analysis of soil enzymes indicated that microbial community were co-limited by C and P during secondary succession. Linear regression of C:N and C:P between the soil nutrient and microbial community showed that soil microbial community maintained stoichiometry homeostasis during plant secondary succession. The threshold elemental ratio revealed that the microbial nutrient metabolisms were co-limited by N and P, particularly P during plant secondary succession. Therefore, microbial communities were co-limited by C, N, and P, particularly C and P during plant secondary succession, and the limitation was mainly associated with soil nutrient status. [ABSTRACT FROM AUTHOR]

Background: Trait based functional and community ecology is en vogue. Most studies, however, ignore phenotypical diversity by characterizing entire species considering only trait means rather than their variability. Phenotypical variability may arise from genotypical differences or from ecological factors (e.g., nutritionally imbalanced diet), and these causes can usually not be separated in natural populations. We used a single genotype from a parthenogenetic model system (the oribatid mite Archegozetes longisetosus Aoki) to exclude genotypical differences. We investigated patterns of dietary (10 different food treatments) induced trait variation by measuring the response of nine different traits (relating to life history, morphology or exocrine gland chemistry). Results: Nutritional quality (approximated by carbon-to-nitrogen ratios) influenced all trait means and their variation. Some traits were more prone to variation than others. Furthermore, the "threshold elemental ratio"- rule of element stoichiometry applied to phenotypic trait variation. Imbalanced food (i.e. food not able to fully meet the nutritional demands of an animal) led to lower trait mean values, but also to a higher variation of traits. Conclusion: Imbalanced food led not only to lower trait value averages, but also to higher trait variability. There was a negative relationship between both parameters, indicating a direct link of both, average trait levels and trait variation to nutritional quality. Hence, variation of trait means may be a predictor for general food quality, and further indicate trade-offs in specific traits an animal must deal with while feeding on imbalanced diets. [ABSTRACT FROM AUTHOR]

Background: Soil microbial communities (SMC) play a central role in the structure and function of desert ecosystems. However, the high variability of annual precipitation could results in the alteration of SMC and related biological processes depending on soil water potential. The nature of the physiological adjustments made by SMC in order to obtain energy and nutrients remains unclear under different soil resource availabilities in desert ecosystems. In order to examine this dynamic, the present study examined the effects of variation in annual precipitation on physiological adjustments by the SMC across two vegetation-soil systems of different soil organic matter input in an oligotrophic desert ecosystem., Methods: We collected soil samples in the Cuatro Ciénegas Basin (Mexico) under two vegetation covers: rosetophylous scrub (RS) and grassland (G), that differ in terms of quantity and quality of organic matter. Collections were conducted during the years 2011, 2012, 2013 and 2014, over which a noticeable variation in the annual precipitation occurred. The ecoenzymatic activity involved in the decomposition of organic matter, and the concentration of dissolved, available and microbial biomass nutrients, were determined and compared between sites and years., Results: In 2011, we observed differences in bacterial taxonomic composition between the two vegetation covers. The lowest values of dissolved, available and microbial nutrients in both cover types were found in 2012. The G soil showed higher values of dissolved and available nutrients in the wet years. Significant positive correlations were detected between precipitation and the ratios Cmic:Nmic and Cmic:Pmic in the RS soil and Cmic:Pmic and Nmic:Pmic in the G soil. The slopes of the regression with Cmic and Nmic were higher in the G soil and lower in the RS soil. Moreover, the SMC under each vegetation cover were co-limited by different nutrients and responded to the sum of water stress and nutrient limitation., Discussion: Soil community within both sites (RS and G) may be vulnerable to drought. However, the community of the site with lower resources (RS) is well adapted to acquire P resources by ecoenzyme upregulation during years with adequate precipitation, suggesting that this community is resilient after drought occurs. Under the Global Climate Change scenarios for desert ecosystems that predict reduced annual precipitation and an increased intensity and frequency of torrential rains and drought events, the soil microbial communities of both sites could be vulnerable to drought through C and P co-limitation and reallocation of resources to physiological acclimatization strategies in order to survive., Competing Interests: Valeria Souza is an Academic Editor for PeerJ.