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Root exudation and its mediated nutrient cycling process driven by nitrogen (N) fertilizer can stimulate the plant availability of various soil nutrients, which is essential for microbial nutrient acquisition. However, the response of soil microbial resource limitations to long-term N fertilizer application rates in greenhouse vegetable systems has rarely been investigated. Therefore, we selected a 15-year greenhouse vegetable system, and investigated how N fertilizer application amount impacts on root carbon and nitrogen exudation rates, microbial resource limitations and microbial carbon use efficiency (CUEST). Four N treatments were determined: high (N3), medium (N2), low (N1), and a control without N fertilization (N0). Compared to the control (N0), the results showed that the root C exudation rates decreased significantly by 42.9 %, 57.3 % and 33.6 %, and the root N exudation rates decreased significantly by 29.7 %, 42.6 %, and 24.1 % under N1, N2, and N3 treatments, respectively. Interactions between fertilizer and plant roots altered microbial C, N, P limitations and CUEST; Microbial C and N/P limitations were positively correlated with root C and N exudation rates, negatively correlated with microbial CUEST. Random Forest analysis revealed that the root C and N exudation rates were key factors for soil microbial resource limitations and microbial CUEST. Through the structural equation model (SEM) analysis, soil NH4+ content had significant direct effects on the root exudation rates after long-term N fertilizer application. An increase in root exudation rates led to enhanced microbial resource limitations in the rhizosphere soils, potentially due to increased competition. This enhancement may reduce microbial carbon use efficiency (CUE), that is, microbial C turnover, thereby reducing soil C sequestration. Overall, this study highlights the critical role of root exudation rates in microbial resource limitations and CUE changes in plant-soil systems, and further improves our understanding of plant-microbial interactions.

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Alpine ecosystems are important terrestrial carbon (C) pools, and microbial decomposers play a key role in litter decomposition. Microbial metabolic limitations in these ecosystems, however, remain unclear. The objectives of this study aim to elucidate the characteristics of microbial nutrient limitation and their C use efficiency (CUE), and to evaluate their response to environmental factors. Five ecological indicators were utilized to assess and compare the degree of microbial elemental homeostasis and the nutrient limitations of the microbial communities among varying stages of litter decomposition (L, F, and H horizon) along an altitudinal gradient (2800, 3000, 3250, and 3500 m) under uniform vegetation (Abies fabri) on Gongga Mountain, eastern Tibetan Plateau. In this study, microorganisms in the litter reached a strictly homeostatic of C content exclusively during the middle stage of litter decomposition (F horizon). Based on the stoichiometry of soil enzymes, we observed that microbial N- and P-limitation increased during litter degradation, but that P-limitation was stronger than N-limitation at the late stages of degradation (H horizon). Furthermore, an increase in microbial CUE corresponded with a reduction in microbial C-limitation. Additionally, redundancy analysis (RDA) based on forward selection further showed that microbial biomass C (MBC) is closely associated with the enzyme activities and their ratios, and MBC was also an important factor in characterizing changes in microbial nutrient limitation and CUE. Our findings suggest that variations in MBC, rather than N- and P-related components, predominantly influence microbial metabolic processes during litter decomposition on Gongga Mountain, eastern Tibetan Plateau.

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Efficient remediation of soil contaminated by polycyclic aromatic hydrocarbons (PAHs) is challenging. To determine whether soil ecoenzyme stoichiometry influences PAH degradation under biostimulation and bioaugmentation, this study initially characterized soil ecoenzyme stoichiometry via a PAH degradation experiment and subsequently designed a validation experiment to answer this question. The results showed that inoculation of PAH degradation consortia ZY-PHE plus vanillate efficiently degraded phenanthrene with a K value of 0.471 (depending on first-order kinetics), followed by treatment with ZY-PHE and control. Ecoenzyme stoichiometry data revealed that the EEAC:N, vector length and angle increased before day five and decreased during the degradation process. In contrast, EEAN:P decreased and then increased. These results indicated that the rapid PAH degradation period induced more C limitation and organic P mineralization. Correlation analysis indicated that the degradation rate K was negatively correlated with vector length, EEAC:P, and EEAN:P, suggesting that C limitation and relatively less efficient P mineralization could inhibit biodegradation. Therefore, incorporating liable carbon and acid phosphatase or soluble P promoted PAH degradation in soils with ZY-PHE. This study provides novel insights into the relationship between soil ecoenzyme stoichiometry and PAH degradation. It is suggested that soil ecoenzyme stoichiometry be evaluated before designing bioremeiation stragtegies for PAH contanminated soils.

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Soil ecoenzymatic stoichiometry reflects the dynamic equilibrium between microorganism's nutrient requirements and resource availability. However, uncertainties persist regarding the key determinants of nutrient restriction in relation to microbial metabolism under varying degrees of warming. Our long-term and multi-level warming field experiment (control treatment, +0.42 °C, +1.50 °C, +2.55 °C) in a typical alpine meadow unveiled a decline in carbon (C)- and nitrogen (N)-acquired enzymes with escalating warming magnitudes, while phosphorus (P)-acquired enzymes displayed an opposite trend. Employing enzymatic stoichiometry modeling, we assessed the nutrient limitations of microbial metabolic activity and found that C and N co-limited microbial metabolic activities in the alpine meadow. Remarkably, high-level warming (+2.55 °C) exacerbated microbe N limitation, but alleviate C limitations. The structural equation modeling further indicated that alterations in soil extracellular enzyme characteristics (SES) were more effectively elucidated by microbial characteristics (microbial biomass C, N, P, and their ratios) rather than by soil nutrients (total nutrient contents and their ratios). However, the microbial control over SES diminished with higher levels of warming magnitude. Overall, our results provided novel evidence that the factors driving microbe metabolic limitation may vary with the degree of warming in Tibet alpine grasslands. Changes in nutrient demand for microorganism's metabolism in response to warming should be considered to improve nutrient management in adapting to different future warming scenarios.

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Deciphering the pivotal components of nutrient metabolism in compost is of paramount importance. To this end, ecoenzymatic stoichiometry, enzyme vector modeling, and statistical analysis were employed to explore the impact of exogenous ore improver on nutrient changes throughout the livestock composting process. The total phosphorus increased from 12.86 to 18.72 g kg-1, accompanied by a marked neutralized pH with ore improver, resulting in the Carbon-, nitrogen-, and phosphorus-related enzyme activities decreases. However, the potential C:P and N:P acquisition activities represented by ln(ßG + CB): ln(ALP) and ln(NAG): ln(ALP), were increased with ore improver addition. Based on the ecoenzymatic stoiometry theory, these changes reflect a decreasing trend in the relative P/N limitation, with pH and total phosphorus as the decisive factors. Our study showed that the practical employment of eco stoichiometry could benefit the manure composting process. Moreover, we should also consider the ecological effects from pH for the waste material utilization in sustainable agriculture.

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Aquatic litter decomposition is highly dependent on contributions and interactions at different trophic levels. The invasion of alien aquatic organisms like the channeled apple snail (Pomacea canaliculata) might lead to changes in the decomposition process through new species interactions in the invaded wetland. However, it is not clear how aquatic macroinvertebrate predators like the Chinese mitten crab (Eriocheir sinensis) will affect the nutrient cycle in freshwater ecosystems in the face of new benthic invasion. We used the litter bag method to explore the top-down effect of crabs on the freshwater nutrient cycle with the help of soil zymography (a technology previously used in terrestrial ecosystems). The results showed significant feeding effects of crabs and snails on lotus leaf litter and cotton strips. Crabs significantly inhibited the intake of lotus litter and cotton strips and the ability to transform the environment of snails by predation. Crabs promoted the decomposition of various litter substrates by affecting the microbial community structure in the sediment. These results suggest that arthropod predators increase the complexity of detrital food webs through direct and indirect interactions, and consequently have an important impact on the material cycle and stability of freshwater ecosystems. This top-down effect makes macrobenthos play a key role in the biological control and engineering construction of freshwater ecosystems.

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The status of plant and microbial nutrient limitation have profound impacts on ecosystem carbon cycle in permafrost areas, which store large amounts of carbon and experience pronounced climatic warming. Despite the long-term standing paradigm assumes that cold ecosystems primarily have nitrogen deficiency, large-scale empirical tests of microbial nutrient limitation are lacking. Here we assessed the potential microbial nutrient limitation across the Tibetan alpine permafrost region, using the combination of enzymatic and elemental stoichiometry, genes abundance and fertilization method. In contrast with the traditional view, the four independent approaches congruently detected widespread microbial nitrogen and phosphorus co-limitation in both the surface soil and deep permafrost deposits, with stronger limitation in the topsoil. Further analysis revealed that soil resources stoichiometry and microbial community composition were the two best predictors of the magnitude of microbial nutrient limitation. High ratio of available soil carbon to nutrient and low fungal/bacterial ratio corresponded to strong microbial nutrient limitation. These findings suggest that warming-induced enhancement in soil nutrient availability could stimulate microbial activity, and probably amplify soil carbon losses from permafrost areas.

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Soil fertility can be increased by returning crop residues to fields due to the cooperative regulation of microbial metabolism of carbon (C) and nutrients. However, the dose-effect of straw on the soil C and nutrient retention and its underlying coupled microbial metabolic processes of C and nutrients remain poorly understood. Here, we conducted a comprehensive study on soil nutrients and stoichiometry, crop nutrient uptake and production, microbial metabolic characteristics and functional attributes using a long-term straw input field experiment. We estimated the microbial metabolic limitations and efficiency of C and nitrogen (N) use (CUE and NUE) via an enzyme-based vector-TER model, biogeochemical-equilibrium model and mass balance equation, respectively. In addition, the absolute abundances of 20 functional genes involved in the N- and P-cycles were quantified by quantitative PCR-based chip technology. As expected, straw input significantly increased C and N stocks, C: nutrients, crop nutrient uptake and growth. However, the C sequestration efficiency decreased by approximately 6.1 %, and the N2O emission rate increased by 0.5-1.0 times with the increase in straw input rate. Interestingly, the microbial metabolism was more limited by P when straw input was <8 t ha-1 but was reversed when straw input was 12 t ha-1. The enhanced nutrient limitation reduced both the CUE and the NUE of microbes and then upregulated genes associated with the hydrolysis of C, the mineralization of N and P, and denitrification, which consequently influenced C and N losses as well as crop growth. This study highlights that soil C and nutrient cycling are strongly regulated by microbial metabolic limitation, suggesting that adding the appropriate limiting nutrients to reduce nutrient imbalances caused by straw input is conducive to maximizing the ecological benefits of straw return.

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Potentially toxic elements (PTEs) pollution causes a great threat to microbial metabolism, which plays a vital role in studying soil nutrient cycling and predicting carbon (C) storage in agroecosystems. However, the responses of microbial metabolism characteristic to heavy metal contamination and the mechanisms through which microbial metabolism mediate nutrient cycling and C dynamics in contaminated soil remain elusive. Here, we performed an incubation experiment over 80 days to investigate the variations in microbial metabolic limitation under various Pb levels (0, 100, 500, 800, 1500, 2000, and 3000 mg Pb kg-1 dry soil) in cropland soil using extracellular enzymatic stoichiometry, and to reveal the impact of Pb stress on soil C storage by associating with microbial metabolic quotients (qCO2) and C use efficiency (CUE). The results showed microbial relative C limitation and phosphorus (P) limitation were observed in Pb-contaminated soils. Pb addition enhanced the microbial relative C limitation by approximately 7.3%, while decreasing the P limitation by approximately 12.3%. Furthermore, Pb addition led to higher qCO2 (from 8.75 to 108 µg C kg-1 MBC-1 d-1) duo to the increase of microbial relative C limitation, suggesting that the more CO2 was released of per unit of microbial biomass C. The increase of microbial relative C limitation reduced CUE (from 0.35 to 0.10) because of the change in microbial metabolism from growth to respiration maintenance under Pb stress. Consequently, the CUE and qCO2 together determined the loss of soil C. Our study reveals that microbial relative C limitation is the dominant driver of soil C loss and provides important knowledge of microbial metabolic limitation regulating soil C turnover in PTEs-contaminated agricultural soils.

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Soil extracellular enzyme activities of microbes to acquire carbon (C), nitrogen (N) and phosphorus (P) exert great roles on soil C sequestration and N, P availability. However, a lack of biochar-induced changes of C, N and P acquisition enzyme activities hinders us from understanding if biochar application will lead to microbial C, N and P limitation based on ecoenzymatic stoichiometry. In this study, through ecoenzymatic stoichiometry, a meta-analysis was conducted to evaluate responses of microbial metabolic limitation to biochar amendment by collecting data of ecoenzymatic activities (EEAs) of the C, N and P acquisition from peer-reviewed papers. The results showed that biochar application increased activities of C, N acquisition enzymes significantly by 9.3 % and 15.1 % on average, respectively. But the influence on P acquisition enzymes activities (Acid, neutral or alkaline phosphatase, abbreviated wholly as PHOS) was not significant. Biochar increased ratio of C acquisition enzymes activities (EC) over P enzymes activities (EP) and ratio of N enzymes activities (EN) over EP, but decreased EC:EN, indicating an increased N limitation or a shift from P limitation to N limitation in microbial metabolism. Enzyme vector analysis showed that soil microbial metabolism was limited by C relative to nutrients (N and P) under biochar amendment according to the overall increased vector length (~1.5 %). Wood biochar caused the strongest microbial C limitation, followed by crop residue biochar as indicated by increased enzyme vector length of 3.6 % and 1.2 % on average, respectively. The stronger microbial C limitation was also found when initial soil total organic carbon (SOC) was <20 g·kg-1. Our results illustrated that available nitrogen and organic carbon should be provided to meet microbial stoichiometric requirements to improve plant productivity, especially in low fertile soils under biochar amendment.

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Microbial nitrogen (N) limitation is a common problem in terrestrial ecosystems. Pig manure, a type of solid waste, is increasingly applied to improve soil N availability in agriculture through inputs of organic matter and inorganic N. Pig manure application also introduces a lot of exogenous microorganisms, which have distinctly different N requirements and metabolic properties, into the resident soil microbial community. However, the impacts of these manure-borne microorganisms on soil N cycling have not been well determined. Here, we investigated effects of manure-borne microorganisms on the N limitation of soil microorganisms using an ecoenzymatic stoichiometry analysis. We monitored microbial communities over a 90-day period in a laboratory-controlled experiment with four treatments: (1) non-sterilized soil mixed with non-sterilized manure (S-M), (2) non-sterilized soil mixed with sterilized manure (S-sM), (3) sterilized soil mixed with non-sterilized manure (sS-M), and (4) non-sterilized soil without manure addition (S, the control). The microbial N limitations were significantly mitigated in both S-M and sS-M. By contrast, the S-sM and S showed high levels of microbial N limitation, likely stemming from differences in the microbial functional composition. We found chitin-degrading bacteria were the dominant copiotrophic manure-borne bacteria associated with N mineralization, and they may improve soil N availability. We further identified several copiotrophic manure-borne bacteria in S-M and sS-M, and their abundances had significantly negative correlation with the level of N limitation and significantly positive correlation with the stoichiometric homeostasis. As these copiotrophic taxa can maintain homeostasis through regulating enzymatic activities, our results indicate that copiotrophic taxa in pig manure contribute to the mitigation of soil microbial N limitation. Our study also highlights the invasiveness capacity of manure-borne microorganisms in soil and evaluates the biotic effects of manure application on soil N cycling.

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Understanding changes in soil enzyme activities and ecoenzymatic stoichiometry is important for assessing soil nutrient availability and microbial nutrient limitation in mountain ecosystems. However, the variations of soil microbial nutrient limitation across elevational gradients and its driving factors in subtropical mountain forests are still unclear. In this study, we measured soil properties, microbial biomass, and enzyme activities related to carbon (C), nitrogen (N), and phosphorus (P) cycling in Pinus taiwanensis forests at different altitudes of Wuyi Mountains. By analyzing the enzyme stoichiometric ratio, vector length (VL), and vector angle (VA), the relative energy and nutrient limitation of soil microorganisms and its key regulatory factors were explored. The results showed that ß-glucosaminidase (BG) activities increased along the elevational gradient, while the activities of ß-N-acetyl glucosaminidase (NAG), leucine aminopeptidase (LAP), acid phosphatase (AcP) and (NAG+LAP)/microbial biomass carbon (MBC) and AcP/MBC showed the opposite trend. Enzyme C/N, enzyme C/P, enzyme N/P, and VL were enhanced with increasing elevation, while VA decreased, indicating a higher degree of microbial P limitation at low elevation and higher C limitation at high elevation. In addition, our results suggested that dissolved organic carbon and microbial biomass phosphorus are critical factors affecting the relative energy and nutrient limitation of soil microorganisms at different elevations. The results would provide a theoretical basis for the responses of soil carbon, nitrogen, and phosphorus availability as well as the relative limitation of microbial energy and nutrition to elevational gradients, and improve our understanding of soil biogeochemical cycle process in subtropical montane forest ecosystems.

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Slow nutrient turnover and destructed soil function were the main factors causing low efficiency in phytoremediation of heavy metal (HM)-contaminated soil. Soil ecoenzymatic stoichiometry can reflect the ability of soil microorganisms to acquire energy and nutrients, and drive nutrient cycling and carbon (C) decomposition in HM-contaminated soil. Therefore, for the first time, we used the enzymatic stoichiometry modeling to examine the microbial nutrient limitation in rhizospheric and bulk soil of different plants (Medicago sativa, Halogeton arachnoideus and Agropyron cristatum) near the Baiyin Copper Mine. Results showed that the main pollutants in this area were Cu, Zn, Cd, and Pb, while Cd and Zn have the greatest contribution according to the analysis of pollution load index (PLI). The activities of soil C-, nitrogen (N)-, and phosphorus (P)-acquiring enzymes in the rhizosphere of plants were significantly greater than that in bulk soil. Moreover, microbial C and P limitations were observed in all plant treatments, while the lower limitation was generally in the rhizosphere compared to bulk soil. The HM stress significantly increased microbial C limitation and decreased microbial P limitation, especially in the rhizospheric soil. The partial least squares path modeling (PLS-PM) further indicated that HM concentration has the greatest effects on microbial P limitation (-0.64). In addition, the highest enzyme activities and the lowest P limitation were observed in the rhizospheric and bulk soil of M. sativa, thereby implying that soil microbial communities under the remediation of M. sativa were steadier and more efficient in terms of their metabolism. These findings are important for the elucidation of the nutrient cycling and microbial metabolism of rhizosphere under phytoremediation, and provide guidance for the restoration of HM-contaminated soil.

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Soil extracellular enzymes plays key roles in ecosystem carbon (C), nitrogen (N), and phosphorus (P) cycling, and are very sensitive to climatic, plant, and edaphic factors. However, the interactive effects of these factors on soil enzyme activities at large spatial scales remain unclear. Here, we investigated the spatial pattern of the activities of five soil hydrolyzing enzymes [ß-D-cellobiohydrolase (CB), ß-1,4-glucosidase (BG), ß-1,4-N-acetyl-glucosaminidase (NAG), L-leucine aminopeptidase (LAP), and acid phosphatase (AP)], and their C:N:P acquisition ratios in relation to plant inputs and edaphic properties across a 600-km climatic gradient in secondary grasslands of subtropical China. The activities of CB, BG, and NAG decreased while that of LAP increased with the increasing mean annual temperature (MAT). The activities of all enzymes did not significantly vary with the mean annual precipitation (MAP). We found that the activities of BG, NAG, and AP were predominately dependent on plant N contents, while the soil LAP activity was tightly related to soil recalcitrant C and N contents. In contrast, the ecoenzymatic C:nutrient (N and P) acquisition ratios increased with increasing MAP and decreasing MAT, primarily due to the increase in plant input at warmer and wetter sites. In addition to climates, plant C inputs, C use efficiency, soil pH, soil organic C, soil C:P, and N:P ratios explained 79% and 72% of the overall variation in ecoenzymatic C:nutrient and P:N acquisition ratios, respectively. The pattern of ecoenzymatic C:N:P acquisition ratios also revealed unexpected N limitation in subtropical grasslands. Overall, our study highlighted the importance of climate in controlling soil biological C, N, and P acquisition activities through its direct and indirect effects on plant inputs and soil edaphic factors, thereby providing useful information for better understanding and predictions of soil C and nutrient cycling in grassland ecosystems at regional scales.

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Soil enzyme activity is an important index to characterize the nutrient requirements and nutrient limitations of soil microorganisms. In this study, Pinus massoniana plantations of different stand ages (9, 17, 26, 34, and 43 a) in mid-subtropical China were taken as the research object; the activities of ß-glucosidase (BG), N-acetyl-ß-glucosaminidase (NAG), leucine amino-peptidase (LAP), acid phosphatase (AP), polyphenol oxidase (POX), and peroxidase (POD) were determined; and soil enzyme stoichiometric ratios were also calculated to investigate the soil microbial nutrient limitations of P. massoniana plantation development. The results showed that the activities of BG, NAG, AP, POX, and POD were enhanced with the increase in stand age, and the activity of LAP was the lowest at 17 a, which showed a significant difference and fluctuated among the neighboring stand ages. The soil enzyme C:N:P stoichiometric ratio was 1:1:0.56, which deviated from the global ecosystem enzyme C:N:P stoichiometric ratio (1:1:1). The enzyme C:N increased, whereas the enzyme N:P decreased, with increasing stand age, and both ratios tended to be stable after 17 a. There was no significant difference in enzyme N:P among different stand ages. The vector length of enzyme stoichiometry was not significantly different among the five stand ages. The vector angles increased with the increase in stand ages and tended to be stable after 17 a of stand age, but the angles were less than 45°. Redundancy analysis (RDA) revealed that soil carbon quality index and pH were the main factors influencing soil enzyme activity and the associated stoichiometric ratio. Our findings indicated that P. massoniana plantation soil microorganisms at different growth stages were all subjected to N limitation, and the N limitation was alleviated with the increase in stand age; however, the P requirement was gradually enhanced. Therefore, the management of P. massoniana plantations should take care to increase nitrogen fertilizer at the early growth stage of P. massoniana, and more phosphorus fertilizers need to be applied with nitrogen at the late growth stage in order to maintain the productivity and sustainable development of P. massoniana plantations.

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Characterized by continuous chemical fertilization, intensive agriculture generally reduces soil ecoenzymatic activities and nutrient mineralization, as well as alters the biomass production and microbial community composition. Soil acidification poses serious threats to the sustainable development of intensive agriculture. However, the mechanism of nutrient cycling and metabolism of soil microorganisms in response to soil acidification in intensive agriculture remains unclear. Herein, we studied the variations in ecoenzymatic stoichiometry of soil ß-glucosidase (BG), cellobiohydrolase (CBH), N-acetylglucosaminidase (NAG) and acid phosphatase (AP) under different land use types and pH gradients of tea garden soils. The results revealed that natural forest and cropland soils had significantly higher BG and CBH activities than tea garden soils. Soil BG and CBH activities displayed significant positive correlations with soil pH, total nitrogen (TN) and phosphorus (TP), while soil NAG activity was significantly associated with nitrate nitrogen, total carbon (TC), TN, carbon: phosphorus (C:P) and nitrogen: phosphorus (N:P) ratios. Soil AP activity showed significant negative associations with pH, TP and C:N ratio, but was significantly positively correlated with TC, TN, C:P and N:P ratios. Enzyme vector model revealed that soil microorganisms are limited by P (enzyme vector angle >45°) regardless of land use types. Compared to natural forest soils, the P limitation of microorganisms in tea garden soils became increasingly serious with a decreasing pH gradient, as indicated by the significant increase in enzyme vector angle. Thus, the overall ecoenzymatic stoichiometry was shifted by soil pH. In summary, higher pH increased BG activity and decreased AP activity, but had no significant effect on NAG activity, suggesting co-limitation of soil microorganisms by C and P in this area. This study provides novel insights into the effect of soil acidification on ecoenzymatic stoichiometry, and also highlights the stoichiometric and energy limitations on the metabolism of soil microorganisms in agricultural ecosystems.

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BACKGROUND: Soil- and plant-produced extracellular enzymes drive nutrient cycling in soils and are assumed to regulate supply and demand for carbon (C) and nutrients within the soil. Thus, agriculture management decisions that alter the balance of plant and supplemental nutrients should directly alter extracellular enzyme activities (EEAs), and EEA stoichiometry in predictable ways. We used a 12-year experiment that varyied three major continuous grain crops (wheat, soybean, and maize), each crossed with mineral fertilizer (WCF, SCF and MCF, respectively) or not fertilized (WC, SC and MC, respectively, as controls). In response, we measured the phospholipid fatty acids (PLFAs), EEAs and their stoichiometry to examine the changes to soil microbial nutrient demand under the continuous cropping of crops, which differed with respect to the input of plant litter and fertilizer. RESULTS: Fertilizer generally decreased soil microbial biomass and enzyme activity compared to non-fertilized soil. According to enzyme stoichiometry, microbial nutrient demand was generally C- and phosphorus (P)-limited, but not nitrogen (N)-limited. However, the degree of microbial resource limitation differed among the three crops. The enzymatic C:N ratio was significantly lower by 13.3% and 26.8%, whereas the enzymatic N:P ratio was significantly higher by 9.9% and 42.4%, in MCF than in WCF and SCF, respectively. The abundances of arbuscular mycorrhizal fungi and aerobic PLFAs were significantly higher in MCF than in WCF and SCF. CONCLUSION: These findings are crucial for characterizing enzymatic activities and their stoichiometries that drive microbial metabolism with respect to understanding soil nutrient cycles and environmental conditions and optimizing practices of agricultural management. © 2021 Society of Chemical Industry.

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Mountain glaciers retreat at an increased rate under global warming, resulting in exposed barren surfaces for primary succession. Soil microbes are an important driver of ecosystem processes. Although variations in soil microbes after deglaciation have been studied extensively, the roles of rhizosphere soil microbes in the biogeochemistry cycle during primary succession are less understood. In this study, Populus purdomii was present throughout the 123-year chronosequence as a representative tree species. We therefore investigated variations in the rhizosphere enzyme activity, microbial community structure, and ecoenzymatic stoichiometry of P. purdomii along Hailuogou Glacier chronosequences. The objective was to determinechanges in rhizosphere enzyme activities and microbial communities, as well as the effects of nutrient limitation on rhizosphere microbes. According to the results, the enzyme activities and microbial group biomass in rhizosphere soil all showed a bimodal trend and were highest at the 43rd or 123rd year, and enzyme activity varied with succession time but not microbial community structure. The rhizosphere soil bacterial community was the dominant community during the 123-year chronosequence. Ecoenzymatic stoichiometry indicated nitrogen restrictions on microbial activity throughout primary succession, with early succession stages (5-15 years) showing greater carbon restriction than late succession stages. Moreover, redundancy and correlation analyses demonstrated that soil microbial phospholipid fatty acid biomass was an important factor for increases in enzyme activities and that enzyme activities in turn played important roles in carbon, nitrogen and phosphorus cycling in rhizosphere soil. Additionally, rhizosphere soil microbial development significantly affected soil organic carbon, total nitrogen and dissolved organic carbon accumulation. Overall, our study links the rhizosphere microbial community and activity to successional chronosequences, providing a deeper understanding of the dynamics of ecosystem succession.

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We measured the activities of six kinds of enzyme, including ß-glucosidase (BG), ß-N-acetyl-glucosaminidase (NAG), leucine aminopeptidase (LAP), acid phosphatase (AP), polyphenol oxidase (POX), peroxidase (POD), as well as enzyme stoichiometric ratios and soil physical and chemical properties at 0-10 and 10-20 cm layers across typical Pinus massoniana plantation, Pinus elliottii plantation and mixed plantation of P. massoniana and Schima superba (broadleaved-conifer mixed plantation) in mid-subtropical China. Key factors driving the variation in soil enzyme activity and stoichiometry among different stand types were investigated. The results showed that the activities of soil BG and LAP were significantly affected by stand type. Soil BG activity at 10-20 cm soil layer was significantly higher in P. elliottii plantation than in P. massoniana plantation, while the activity of LAP was highest in the P. massoniana plantation. Soil BG/(NAG+LAP) and BG/AP at 10-20 cm layer of P. elliottii plantation were significantly higher than those of P. massoniana plantation, while (NAG+LAP)/AP of P. massoniana plantation was significantly higher than those of P. elliottii plantation and mixed plantation. The vector length of enzyme stoichiometry at 10-20 cm soil layer was significantly different among stand type, with an order of P. elliottii plantation > broadleaved-conifer mixed plantation > P. massoniana. The vector angles of enzyme stoichiometry in the three plantations were greater than 45°, with the vector angle in the P. elliottii plantation at 10-20 cm soil layer being significantly greater than that of the P. massoniana plantation. Results from redundancy analysis showed that soil carbon quality index and the ratio of soil organic carbon to total phosphorus (C/P), soil water content and C/P were the key factors affecting soil enzyme activity and stoichiometry at 0-10 and 10-20 cm soil layers, respectively. The quantity and quality of soil carbon and phosphorus, and soil water content played a key role in regulating nutrient cycling in mid-subtropical plantation ecosystem.

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Knowledge about resource limitation to soil microbes is crucial for understanding ecosystem functions and processes, and for predicting ecosystem responses to global changes as well. Karst ecosystems are widespread in the world, and play a key role in regulating the global climate, however, the patterns of and mechanisms underlying microbial resource limitation in karst ecosystems remain poorly known. Here we investigated the microbial resource limitation in a karst region, by selecting four main land-use types, i.e. cropland, grassland, shrubland and secondary forest, in areas underlain by two lithology types, i.e. dolomite and limestone, in southwest China. Ecoenzymatic stoichiometry was used as an indicator of microbial resource limitation. Overall, soil microbes in karst ecosystems were more limited by carbon and phosphorus, rather than by nitrogen. Further analyses revealed that the patterns of carbon and phosphorus limitation were different among land-use or lithology types. Microbial carbon limitation was greatest in cropland and forest but lowest in grassland, and was greater under dolomite than under limestone. Microbial phosphorus limitation decreased from secondary forest to cropland under dolomite areas, but showed no difference among ecosystem types under limestone areas, indicating that lithology controls the pattern of microbial phosphorus limitation along the post-agriculture succession. Our study describes a general pattern of microbial resource limitation in karst ecosystems, and we suggest that lithology may provide a new mechanism for explaining the variations of microbial resource limitation along the post-agriculture succession in different regions.

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