Microbial immobilisation of C rhizodeposits in rhizosphere and root-free soil under continuous C labelling of oats

A greenhouse experiment was conducted by growing oats (Avenasativa L.) in a continuously ¹³CO₂ labeled atmosphere. The allocation of ¹³C-labeled photosynthates in plants, microbial biomass in rhizosphere and root-free soil, pools of soil organic C, and CO₂ emissions were examined over the plant's life cycle. To isolate rhizosphere from root-free soil, plant seedlings were placed into bags made of nylon monofilament screen tissue (16 μm mesh) filled with soil. Two peaks of ¹³C in rhizosphere pools of microbial biomass and dissolved organic carbon (DOC), as well as in CO₂ emissions at the earing and ripeness stages were revealed. These ¹³C maxima corresponded to: (i) the end of rapid root growth and (ii) beginning of root decomposition, respectively. The δ¹³C values of microbial biomass were higher than those of DOC and of soil organic matter (SOM). The microbial biomass C accounted for up to 56 and 39% of ¹³C recovered in the rhizosphere and root-free soil, respectively. Between 4 and 28% of ¹³C assimilated was recovered in the root-free soil. Depending on the phenological stage, the contribution of root-derived C to total CO₂ emission from soil varied from 61 to 92% of total CO₂ evolved, including 4-23% attributed to rhizomicrobial respiration. While 81-91% of C substrates used for microbial growth in the root-free soil and rhizosphere came from SOM, the remaining 9-19% of C substrates utilized by the microbial biomass was attributable to rhizodeposition. The use of continuous isotopic labelling and physical separation of root-free and rhizosphere soil, combined with natural ¹³C abundance were effective in gaining new insight on soil and rhizosphere C-cycling.     

Carbon fluxes in soil food webs of increasing complexity revealed by C labelling and C natural abundance

Soil food webs are mainly based on three primary carbon (C) sources: root exudates, litter, and recalcitrant soil organic matter (SOM). These C sources vary in their availability and accessibility to soil organisms, which could lead to different pathways in soil food webs. The presence of three C isotopes (¹²C, ¹³C and ¹⁴C) offers an unique opportunity to investigate all three C sources simultaneously. In a microcosm experiment we studied the effect of food web complexity on the utilization of the three carbon sources. We choose an incomplete three factorial design with (i) living plants, (ii) litter and (iii) food web complexity. The most complex food web consisted of autochthonous microorganisms, nematodes, collembola, predatory mites, endogeic and anecic earthworms. We traced C from all three sources in soil, in CO₂ efflux and in individual organism groups by using maize grown on soil developed under C₃ vegetation and application of ¹⁴C labelled ryegrass shoots as a litter layer. The presence of living plants had a much greater effect on C pathways than food web complexity. Litter decomposition, measured as ¹⁴CO₂ efflux, was decreased in the presence of living plants from 71% to 33%. However, living plants increased the incorporation of litter C into microbial biomass and arrested carbon in the litter layer and in the upper soil layer. The only significant effect of food web complexity was on the litter C distribution in the soil layers. In treatments with fungivorous microarthropods (Collembola) the incorporation of litter carbon into mineral soil was reduced. Root exudates as C source were passed through rhizosphere microorganisms to the predator level (at least to the third trophic level). We conclude that living plants strongly affected C flows, directly by being a source of additional C, and indirectly by modifying the existing C flows within the food web including CO₂ efflux from the soil and litter decomposition.     

Carbon partitioning in ectomycorrhizal Scots pine seedlings

The complete carbon budget and the turnover rate of assimilated carbon of ectomycorrhizal Scots pine seedlings growing on natural humus were determined in microcosm conditions. The main aim was to improve understanding of the partitioning of the assimilated carbohydrates within seedlings associated with multiple ectomycorrhizal fungi, and to discover carbon dynamics of the mycorrhizosphere.Plant photosynthesis and below-ground respiration were measured in order to obtain the actual carbon assimilation and respiration rates at the time of measurements. Soon after the photosynthesis and respiration rate measurements the seedlings were pulse-labeled with ¹⁴CO₂ to follow carbon allocation to different plant, fungal and soil compartments and rhizosphere respiration. Long-term carbon allocation during the entire life span of the seedlings was estimated by measuring plant and mycorrhizal root-tip biomass. The ectomycorrhizal community was analyzed using morphotyping and ITS-sequencing.The ¹⁴C label was detected in rhizosphere respiration after 12 h and it peaked between 36 and 60 h after labeling. More than half of the assimilated carbon was allocated below-ground as biomass or respiration and higher mycorrhizal biomass increased the below-ground carbon turnover. The presence of Suillus variegatus affected the plant carbon balance in several ways. When S. variegatus was present, the below-ground respiration increased and this carbon loss was compensated by higher photosynthetic activity. Other fungal species did not differ between each other in their effects on carbon balance. Our findings indicate that some root-associated mycorrhizal fungal symbionts can significantly alter plant CO₂ exchange, biomass distribution, and the allocation of recently photosynthesized plant-derived carbon.     

Contribution of exudates,arbuscular mycorrhizal fungi and litter depositions to the rhizosphere priming effect induced by grassland species

The presence of plants induces strong accelerations in soil organic matter (SOM) mineralization by stimulating soil microbial activity – a phenomenon known as the rhizosphere priming effect (RPE). The RPE could be induced by several mechanisms including root exudates, arbuscular mycorrhizal fungi (AMF) and root litter. However the contribution of each of these to rhizosphere priming is unknown due to the complexity involved in studying rhizospheric processes. In order to determine the role of each of these mechanisms, we incubated soils enclosed in nylon meshes that were permeable to exudates, or exudates & AMF or exudates, AMF and roots under three grassland plant species grown on sand. Plants were continuously labeled with ¹³C depleted CO₂ that allowed distinguishing plant-derived CO₂ from soil-derived CO₂. We show that root exudation was the main way by which plants induced RPE (58–96% of total RPE) followed by root litter. AMF did not contribute to rhizosphere priming under the two species that were significantly colonized by them i.e. Poa trivialis and Trifolium repens. Root exudates and root litter differed with respect to their mechanism of inducing RPE. Exudates induced RPE without increasing microbial biomass whereas root litter increased microbial biomass and raised the RPE mediating saprophytic fungi. The RPE efficiency (RPE/unit plant-C assimilated into microbes) was 3–7 times higher for exudates than for root litter. This efficiency of exudates is explained by a microbial allocation of fresh carbon to mineralization activity rather than to growth. These results suggest that root exudation is the main way by which plants stimulated mineralization of soil organic matter. Moreover, the plants through their exudates not only provide energy to soil microorganisms but also seem to control the way the energy is used in order to maximize soil organic matter mineralization and drive their own nutrient supply.     

Production of root-derived material and associated microbial growth in soil at different nutrient levels

Summary Maize plants were grown for 42 days in a sandy soil at two different mineral nutrient levels, in an atmosphere containing ¹⁴CO₂. The ¹⁴C and total carbon contents of shoots, roots, soil and soil microbial biomass were measured 28, 35 and 42 days after germination. Relative growth rates of shoots and roots decreased after 35 days at the lower nutrient level, but were relatively constant at the higher nutrient level. In the former treatment, 2% of the total ¹⁴C fixed was retained as a residue in soil at all harvests while at the higher nutrient level up to 4% was retained after 42 days. Incorporation of ¹⁴C into the soil microbial biomass was close to its maximum after 35 days at the lower nutrient level, but continued to increase at the higher level. Generally a good agreement existed between microbial biomass, ¹⁴C contents and numbers of fluorescent pseudomonads in the rhizosphere. Numbers of fluorescent pseudomonads in the rhizosphere were maximal after 35 days at the lower nutrient level and continued to increase at the higher nutrient level. The proportions of the residual ¹⁴C in soil, incorporated in the soil microbial biomass, were 28% to 41% at the lower nutrient level and 20%6 – 30% at the higher nutrient level. From the lower nutrient soil 18%6 – 52%6 of the residual soil ¹⁴C could be extracted with 0.5 N K₂SO₄, versus 14%6 – 16% from the higher nutrient soil.Microbial growth in the rhizosphere seemed directly affected by the depletion of mineral nutrients while plant growth and the related production of root-derived materials continued.     

Atmospheric CO2 enrichment and nutrient additions to planted soil increase mineralisation of soil organic matter, but do not alter microbial utilisation of plant- and soil C-sources

Plants link atmospheric and soil carbon pools through CO₂ fixation, carbon translocation, respiration and rhizodeposition. Within soil, microbial communities both mediate carbon-sequestration and return to the atmosphere through respiration. The balance of microbial use of plant-derived and soil organic matter (SOM) carbon sources and the influence of plant-derived inputs on microbial activity are key determinants of soil carbon-balance, but are difficult to quantify. In this study we applied continuous ¹³C-labelling to soil-grown Lolium perenne, imposing atmospheric CO₂ concentrations and nutrient additions as experimental treatments. The relative use of plant- and SOM-carbon by microbial communities was quantified by compound-specific ¹³C-analysis of phospholipid fatty acids (PLFAs). An isotopic mass-balance approach was applied to partition the substrate sources to soil respiration (i.e. plant- and SOM-derived), allowing direct quantification of SOM-mineralisation. Increased CO₂ concentration and nutrient amendment each increased plant growth and rhizodeposition, but did not greatly alter microbial substrate use in soil. However, the increased root growth and rhizosphere volume with elevated CO₂ and nutrient amendment resulted in increased rates of SOM-mineralisation per experimental unit. As rhizosphere microbial communities utilise both plant- and SOM C-sources, the results demonstrate that plant-induced priming of SOM-mineralisation can be driven by factors increasing plant growth. That the balance of microbial C-use was not affected on a specific basis may suggest that the treatments did not affect soil C-balance in this study.     

Additions of maize root mucilage to soil changed the structure of the bacterial community

The organic compounds released from roots (rhizodeposits) stimulate the growth of the rhizosphere microbial community. They may be responsible for the differences in the structure of the microbial communities commonly observed between the rhizosphere and the bulk soil. Rhizodeposits consists of a broad range of compounds including root mucilage. The aim of this study was to investigate if additions of maize root mucilage, at a rate of 70 μg C g⁻¹ day⁻¹ for 15 days, to an agricultural soil could affect the structure of the bacterial community. Mucilage additions moderately increased microbial C (+23% increase relative to control), which suggests that the turnover rate of microorganisms consuming this substrate was high. Consistent with this, the number of cultivable bacteria was enhanced by +450%. Catabolic (Biolog^® GN2) and 16S-23S intergenic spacer fingerprints exhibited significant differences between control and mucilage treatments. These data indicate that mucilage can affect both the metabolic and genetic structure of the bacterial community as shown by a greater catabolic potential for carbohydrates. We concluded that mucilage is likely to significantly contribute to differences in the structure of the bacterial communities present in the rhizosphere compared to the bulk soil.     

水稻根际和周围土壤中微生物功能基因丰度与碳、氮状况及其通量之间的关系

Rapid nitrogen（N） transformations and losses occur in the rice rhizosphere through root uptake and microbial activities. However,the relationships between rice roots and rhizosphere microbes for N utilization are still unclear. We analyzed different N forms（NH＋4,NO-3, and dissolved organic N）, microbial biomass N and C, dissolved organic C, CH4 and N2O emissions, and abundance of microbial functional genes in both rhizosphere and bulk soils after 37-d rice growth in a greenhouse pot experiment. Results showed that the dissolved organic C was significantly higher in the rhizosphere soil than in the non-rhizosphere bulk soil, but microbial biomass C showed no significant difference. The concentrations of NH＋4, dissolved organic N, and microbial biomass N in the rhizosphere soil were significantly lower than those of the bulk soil, whereas NO-3in the rhizosphere soil was comparable to that in the bulk soil. The CH4 and N2O fluxes from the rhizosphere soil were much higher than those from the bulk soil. Real-time polymerase chain reaction analysis showed that the abundance of seven selected genes, bacterial and archaeal 16 S rRNA genes, amoA genes of ammonia-oxidizing archaea and ammonia-oxidizing bacteria, nosZ gene, mcrA gene, and pmoA gene, was lower in the rhizosphere soil than in the bulk soil, which is contrary to the results of previous studies. The lower concentration of N in the rhizosphere soil indicated that the competition for N in the rhizosphere soil was very strong, thus having a negative effect on the numbers of microbes. We concluded that when N was limiting, the growth of rhizosphere microorganisms depended on their competitive abilities with rice roots for N.     

Identification of Metabolically Active Rhizosphere Microorganisms by Stable Isotopic Probing of PLFA in Switchgrass

Root-derived rhizodeposits of recent photosynthetic carbon (C) are the foremost source of energy for microbial growth and development in rhizosphere soil. A substantial amount of photosynthesized C by the plants is translocated to belowground and is released as root exudates that influence the structure and function of soil microbial communities with potential inference in nutrient and C cycling in the ecosystem. We applied the ¹³C pulse chase labeling technique to evaluate the incorporation of rhizodeposit-C into the phospholipid fatty acids (PLFAs) in the bulk and rhizosphere soils of switchgrass (Panicum virgatum L.). Soil samples of bulk and rhizosphere were taken at 1, 5, 10 and 20 days after labeling and analyzed for ¹³C enrichment in the microbial PLFAs. Temporal differences of ¹³C enrichment in PLFAs were more prominent than spatial differences. Among the microbial PLFA biomarkers, fungi and Gram-negative (GM-ve) bacterial PLFAs showed rapid enrichment with ¹³C compared to Gram-positive (GM+ve) and actinomycetes in rhizosphere soil. The ¹³C enrichment of actinomycetes biomarker PLFA significantly increased along with sampling time in both soils. PLFAs indicative to fungi, GM-ve and GM+ve showed a significant decrease in ¹³C enrichment over sampling time in the rhizosphere, but a decrease was also observed in GM-ve (16:1ω5c) and fungal biomarker PLFAs in the bulk soil. The relative ¹³C concentration in fungal PLFA decreased on day 10, whereas those of GM-ve increased on day 5 and GM+ve remained constant in the rhizosphere soil. However, the relative ¹³C concentrations of GM-ve and GM+ve increased on days 5 and 10, respectively, and those of fungal remain constant in the bulk soil. The present study demonstrates the usefulness of ¹³C pulse chase labeling together with PLFA analysis to evaluate the active involvement of microbial community groups for utilizing rhizodeposit-C.     

Variable use of plant- and soil-derived carbon by microorganisms in agricultural soils

In this study we used compound specific ¹³C and ¹⁴C isotopic signatures to determine the degree to which recent plant material and older soil organic matter (SOM) served as carbon substrates for microorganisms in soils. We determined the degree to which plant-derived carbon was used as a substrate by comparison of the ¹³C content of microbial phospholipid fatty acids (PLFA) from soils of two sites that had undergone a vegetation change from C3 to C4 plants in the past 20-30 years. The importance of much older SOM as a substrate was determined by comparison of the radiocarbon content of PLFA from soils of two sites that had different ¹⁴C concentrations of SOM.The ¹³C shift in PLFA from the two sites that had experienced different vegetation history indicated that 40-90% of the PLFA carbon had been fixed since the vegetation change took place. Thus PLFA were more enriched in ¹³C from the new C4 vegetation than it was observed for bulk SOM indicating recent plant material as preferentially used substrate for soil microorganisms. The largest ¹³C shift of PLFA was observed in the soil that had high ¹⁴C concentrations of bulk SOM. These results reinforce that organic carbon in this soil for the most part cycles rapidly. The degree to which SOM is incorporated into microbial PLFA was determined by the difference in ¹⁴C concentration of PLFA derived from two soils one with high ¹⁴C concentrations of bulk SOM and one with low. These results showed that 0-40% of SOM carbon is used as substrate for soil microorganisms. Furthermore a different substrate usage was identified for different microorganisms. Gram-negative bacteria were found to prefer recent plant material as microbial carbon source while Gram-positive bacteria use substantial amounts of SOM carbon. This was indicated by ¹³C as well as ¹⁴C signatures of their PLFA. Our results find evidence to support ‘priming’ in that PLFA indicative of Gram-negative bacteria associated with roots contain both plant- and SOM-derived C. Most interestingly, we find PLFA indicative of archeobacteria (methanothrophs) that may indicate the use of other carbon sources than plant material and SOM to a substantial amount suggesting that inert or slow carbon pools are not essential to explain carbon dynamics in soil.     

Effect of heavy metals contamination on root-derived and organic matter-derived CO2 efflux from soil planted with Zea mays

The CO₂ efflux from loamy Haplic Luvisol and heavy metal (HM) uptake by Zea mays L. were studied under increased HM contamination: Cd, Cu, and Ni up to 20, 1000, and 2500 mg kg⁻¹ soil, respectively. Split-root system with contrasting HM concentrations in both soil halves was used to investigate root-mediated HM translocation in uncontaminated soil zones. To separate root-derived and soil organic matter (SOM)-derived CO₂ efflux from soil, ¹⁴CO₂ pulse labeling of 15-, 25-, and 35-days-old plants was applied. The CO₂ evolution from the bare soil was 10.6 μg C–CO₂ d⁻¹ g⁻¹ (32 kg C–CO₂ d⁻¹ ha⁻¹) and was not affected by HM (except 2500 mg Ni kg⁻¹). The average CO₂ efflux from the soil with maize was about two times higher and amounted for about 22.0 μg C–CO₂ d⁻¹ g⁻¹. Portion of assimilates respired in the rhizosphere decreased with plant development from 6.0 to 7.0% of assimilated C for 25-days-old Zea mays to 0.4–2.0% for 45-days-old maize. The effect of the HM on root-derived ¹⁴CO₂ efflux increased with rising HM content in the following order: Cd < Cu < Ni. In Cu and Ni contaminated soils, shoot and root dry matter decreased to 70% and to 50% of the uncontaminated control, respectively. Plants contained much more HM in the roots than in the shoots. A split-root system with contrasting HM concentrations allowed to trace transport of mobile forms of HM by roots from contaminated soil half into the uncontaminated soil half. The portion of mobile HM forms in the soil (1 M NH₄NO₃ extract) increased with contamination and amounted to 9–16%, 2–6% and 1.5–3.5% for Cd, Cu, and Ni, respectively. Corresponding values for the easily available HM (1 M NH₄OAc extract) were 22–52%, 1–20% and 5–8.5%. Heavy metal availability for plants decreased in the following order: Cd > Cu ≥ Ni. No increase of HM availability in the soil was found after maize cultivation.     

Distribution and turnover of recently fixed photosynthate in ryegrass rhizospheres

The cycling of root-deposited photosynthate (rhizodeposition) through the soil microbial biomass can have profound influences on plant nutrient availability. Currently, our understanding of microbial dynamics associated with rhizosphere carbon (C) flow is limited. We used a ¹³C pulse-chase labeling procedure to examine the flow of photosynthetically fixed ¹³C into the microbial biomass of the bulk and rhizosphere soils of greenhouse-grown annual ryegrass (Lolium multiflorum Lam.). To assess the temporal dynamics of rhizosphere C flow through the microbial biomass, plants were labeled either during the transition between active root growth and rapid shoot growth (Labeling Period 1), or nine days later during the rapid shoot growth stage (Labeling Period 2). Although the distribution of ¹³C in the plant/soil system was similar between the two labeling periods, microbial cycling of rhizodeposition differed between labeling periods. Within 24 h of labeling, more than 10% of the ¹³C retained in the plant/soil system resided in the soil, most of which had already been incorporated into the microbial biomass. From day 1 to day 8, the proportion of ¹³C in soil as microbial biomass declined from about 90 to 35% in rhizosphere soil and from about 80 to 30% in bulk soil. Turnover of ¹³C through the microbial biomass was faster in rhizosphere soil than in bulk soil, and faster in Labeling Period 1 than Labeling Period 2. Our results demonstrate the effectiveness of using ¹³C labeling to examine microbial dynamics and fate of C associated with cycling of rhizodeposition from plants at different phenological stages of growth.     

Carbon cycling in subarctic tundra; seasonal variation in ecosystem partitioning based on in situ C pulse-labelling

Carbon assimilation and allocation were studied in a tundra ecosystem in northern Scandinavia. Seasonal variation in the below-ground carbon allocation to dissolved organic carbon (DOC), coarse-, fine-, and hair roots was investigated using in situ ¹⁴C pulse-labelling, adding 2-3 MBq ¹⁴CO₂ dm⁻² to the above-ground vegetation. Combining the allocation data with regression models of the seasonal carbon flux made it possible to estimate a temporally explicit ecosystem carbon allocation budget.The ecosystem was a net source of CO₂, losing on average 0.97 g C m⁻² d⁻¹ to the atmosphere, with little variation through the season. There was, however, significant temporal variation in partitioning of recently assimilated carbon. Allocation to below-ground compartments over 32 days following labelling increased from 18% in June to 55% in September. Above-ground allocation showed the opposite trend. Hair roots and DOC were strong sinks in the autumn. Transport of newly assimilated carbon occurred rapidly throughout the season, ¹⁴C appearing in all sampled pools within 4 h of labelling.The seasonal variation in carbon partitioning observed in this study has implications for the residence time of assimilated carbon in the ecosystem. A relatively greater allocation to rapidly decomposing pools, such as hair roots and DOC, would tend to reduce incorporation into woody tissue, increasing the overall rate of carbon cycling and decreasing ecosystem storage. The results of this study will be of value for building and validating mechanistic models of ecosystem carbon flow in tundra and subarctic ecosystems.     

Effects of food quality, starvation and life stage on stable isotope fractionation in Collembola

Naturally occurring stable isotopes of carbon and nitrogen are powerful tools to investigate food webs, where the ratio of ¹⁵N/¹⁴N is used to assign trophic levels and of ¹³C/¹²C to determine the food source. A shift in δ¹⁵N value of 3‰ is generally suggested as mean difference between two trophic levels, whereas the carbon isotope composition of a consumer is assumed to reflect the signal of its diet. This study investigates the effects of food quality, starvation and life stage on the stable isotope fractionation in fungal feeding Collembola. The fractionation of nitrogen was strongly affected by food quality, i.e. the C/N ratio of the fungal diet. Collembola showed enrichment in the heavier isotope with increasing N concentration of the food source. Δ¹⁵N varied between 2.4‰, which assigns a shift in one trophic level, and 6.3‰, suggesting a shift in two trophic levels. Starvation up to 4 weeks resulted in an increase in the total δ¹⁵N value from 2.8‰ to 4.0‰. Different life stages significantly affected the isotope discrimination by Collembola with juveniles showing a stronger enrichment (Δ¹⁵N=4.9‰) compared to adults (Δ¹⁵N=3.5‰). Δ¹³C varied between −2.1‰ and −3.3‰ depending on the food quality, mainly due to compensational feeding on low quality diet. During starvation δ¹³C value decreased by 1.1‰, whereas the life stage of Collembola had no significant effect on isotopic ratios. The results indicate that the food resource and the physiological status of the consumer have important impact on stable isotope discrimination. They may cause differences in fractionation rate comparable to trophic level shifts, a fact to consider when analysing food web structure.     

菊芋根系分泌物改良滨海盐土的微生物机制

[目的] 探究菊芋在滨海盐土改良过程中的作用机制,分析菊芋和碱蓬根系分泌物的组分差异,明确土壤微生态环境的变化规律,进一步为盐土改良提供理论依据。[方法] 以种植菊芋和自然碱蓬植被为样地,对菊芋和碱蓬的根系分泌物进行对比分析,研究在根系分泌物作用下土壤微生物数量,微生物量碳氮,微生物群落结构以及土壤酶活性的变化,从而系统地阐明根系分泌物介导下盐土改良的微生物机制。[结果] 菊芋根际土壤中含有果糖（2.343×10^-3 g/kg）、葡萄糖（4.235×10^-3 g/kg）、蔗糖（2.670×10^-3 g/kg）,分别是碱蓬根际土壤的9.28,1.52和2.43倍。而菊芋根际与非根际中的果糖含量存在显著性差异（p<0.05）,其根际中含量为非根际的12.02倍。菊芋土壤还含有低聚果糖（蔗果三糖、蔗果四糖和蔗果五糖）,而碱蓬土壤中未检测出低聚果糖。除糖类外,菊芋根系分泌物还含有烷烃、酚、醛、酯、有机酸、醇、酮、酰胺,其组分较碱蓬土壤更为复杂且某些组分为菊芋特有〔1-氯—十八烷、正十六烷酸、2-甲基-Z-4-十四碳烯、十二酮、（Z）-9-十八碳酰胺、苯丙酸十六烷基酯等〕。功能性根系分泌物（如低聚果糖、果糖、十六烷、十八烷酸等）为根际微生物提供碳源、氮源和营养元素的同时,使菊芋根际土壤中微生物数量显著增加（p<0.05）,土壤微生物量碳、氮显著高于碱蓬土壤（p<0.05）,其值分别是碱蓬土壤的1.95和1.6倍,且菊芋根际的微生物量碳、氮约为非根际的1.69和1.50倍,优势菌群（变形菌门、放线菌门、绿弯菌门、酸杆菌门）所占比重达到90%,土壤有益菌群（Actinobacteria和Acidobacteria）的相对丰度显著增加（p<0.05）,土壤生物活性提升。此外,菊芋根际特有的分泌物（十六烷、烯醛等）,抑制了病原菌的生长,优化了微生物群落结构。除过氧化氢酶外,土壤脲酶、蔗糖酶和碱性磷酸活性显著提高（p<0.05）,其活性分别是碱蓬土壤的1.83,1.88和3.30倍。[结论] 种植菊芋后,通过根际分泌物介导,改善土壤微生物群落结构与功能,增加土壤酶活性,使土壤生物活力得以整体提升,与原生植被碱蓬相比,降低了土壤含盐量,起到了改良盐土的作用。     

Rhizospheric influence on soil respiration and decomposition in a temperate Norway spruce stand

Assessments of terrestrial carbon fluxes require a thorough understanding of links between primary production, soil respiration and carbon loss through drainage. In this study, stem girdling was used to terminate autotrophic soil respiration including rhizosphere respiration and root exudation in a temperate Norway spruce stand. Rates of soil respiration and dissolved organic carbon (DOC) formation were measured in the second year after girdling, comparing an intact plant-rhizosphere continuum with an exclusive decomposer system. The molecular and isotopic composition of DOC in the soil solution was analysed with a coupled Py-GC/MS-C-IRMS system to distinguish between the carbon sources of dissolved carbon. Pyrolysis products were grouped according to their precursor origins: polysaccharides, proteins or of mixed origin (mainly derivates of lignins and proteins). When dead roots became available for decomposition, rates of heterotrophic soil respiration in girdling plots peaked at 6.5 μmol m⁻² s⁻¹, comparable to peak rates of total soil respiration (autotrophic and heterotrophic) in control plots, 6.1 μmol m⁻² s⁻¹. A significant response of soil respiration to temperature was found in control plots only, showing that an unlimiting supply of organic substrates for microbial respiration may mask any temperature effects. The enhanced decomposition in girdled plots was further supported by the isotopic composition of DOC in soil solution; all three precursor groups became isotopically enriched as the growing season progressed (polysaccharides by 2.3‰, proteins by 1.9‰, mixed origin group by 2.2‰). This indicates a trophic level shift due to incorporation of organic substrate into the microbial food chain. In the control plots’ mixed origin fraction, the isotopic composition changed over time from a signature resembling that of lignin (−28.9‰) to one similar of the protein fraction (−25.7‰). Significant temporal changes of structural DOC composition occurred in the girdling plots only. These results suggest that changes in the microbial community and in decomposition rates occurred in both girdled and control plots in the following ways: (i) increased substrate availability (dead roots) gave rise to generally enhanced performance of the decomposer community in girdled plots, (ii) root-derived exudates probably contributed to enhanced decomposition of recalcitrant lignin in the control plots and (iii) the structural composition of DOC seemed to be more a result of decomposition than of plant root exudation in all plots.     

Atmospheric nitrate deposition and the microbial degradation of cellobiose and vanillin in a northern hardwood forest

Human activity has increased the amount of N entering terrestrial ecosystems from atmospheric NO₃⁻ deposition. High levels of inorganic N are known to suppress the expression of phenol oxidase, an important lignin-degrading enzyme produced by white-rot fungi. We hypothesized that chronic NO₃⁻ additions would decrease the flow of C through the heterotrophic soil food web by inhibiting phenol oxidase and the depolymerization of lignocellulose. This would likely reduce the availability of C from lignocellulose for metabolism by the microbial community. We tested this hypothesis in a mature northern hardwood forest in northern Michigan, which has received experimental atmospheric N deposition (30 kg NO₃⁻-N ha⁻¹ y⁻¹) for nine years. In a laboratory study, we amended soils with ¹³C-labeled vanillin, a monophenolic product of lignin depolymerization, and ¹³C-labeled cellobiose, a disaccharide product of cellulose degradation. We then traced the flow of ¹³C through the microbial community and into soil organic carbon (SOC), dissolved organic carbon (DOC), and microbial respiration. We simultaneously measured the activity of enzymes responsible for lignin (phenol oxidase and peroxidase) and cellobiose (β-glucosidase) degradation. Nitrogen deposition reduced phenol oxidase activity by 83% and peroxidase activity by 74% when compared to control soils. In addition, soil C increased by 76%, whereas microbial biomass decreased by 68% in NO₃⁻ amended soils. ¹³C cellobiose in bacterial or fungal PLFAs was unaffected by NO₃⁻ deposition; however, the incorporation of ¹³C vanillin in fungal PLFAs extracted from NO₃⁻ amended soil was 82% higher than in the control treatment. The recovery of ¹³C vanillin and ¹³C cellobiose in SOC, DOC, microbial biomass, and respiration was not different between control and NO₃⁻ amended treatments. Chronic NO₃⁻ deposition has stemmed the flow of C through the heterotrophic soil food web by inhibiting the activity of ligninolytic enzymes, but it increased the assimilation of vanillin into fungal PLFAs.     

Rhizodeposition-induced decomposition increases N availability to wild and cultivated wheat genotypes under elevated CO2

Elevated CO₂ may increase nutrient availability in the rhizosphere by stimulating N release from recalcitrant soil organic matter (SOM) pools through enhanced rhizodeposition. We aimed to elucidate how CO₂-induced increases in rhizodeposition affect N release from recalcitrant SOM, and how wild versus cultivated genotypes of wheat mediated differential responses in soil N cycling under elevated CO₂. To quantify root-derived soil carbon (C) input and release of N from stable SOM pools, plants were grown for 1 month in microcosms, exposed to ¹³C labeling at ambient (392 μmol mol⁻¹) and elevated (792 μmol mol⁻¹) CO₂ concentrations, in soil containing ¹⁵N predominantly incorporated into recalcitrant SOM pools. Decomposition of stable soil C increased by 43%, root-derived soil C increased by 59%, and microbial-¹³C was enhanced by 50% under elevated compared to ambient CO₂. Concurrently, plant ¹⁵N uptake increased (+7%) under elevated CO₂ while ¹⁵N contents in the microbial biomass and mineral N pool decreased. Wild genotypes allocated more C to their roots, while cultivated genotypes allocated more C to their shoots under ambient and elevated CO₂. This led to increased stable C decomposition, but not to increased N acquisition for the wild genotypes. Data suggest that increased rhizodeposition under elevated CO₂ can stimulate mineralization of N from recalcitrant SOM pools and that contrasting C allocation patterns cannot fully explain plant mediated differential responses in soil N cycling to elevated CO₂.     

C fractionation at the root-microorganisms-soil interface: A review and outlook for partitioning studies

Natural variations of the ¹³C/¹²C ratio have been frequently used over the last three decades to trace C sources and fluxes between plants, microorganisms, and soil. Many of these studies have used the natural-¹³C-labelling approach, i.e. natural δ¹³C variation after C₃-C₄ vegetation changes. In this review, we focus on ¹³C fractionation in main processes at the interface between roots, microorganisms, and soil: root respiration, microbial respiration, formation of dissolved organic carbon, as well as microbial uptake and utilization of soil organic matter (SOM). Based on literature data and our own studies, we estimated that, on average, the roots of C₃ and C₄ plants are ¹³C enriched compared to shoots by +1.2 ± 0.6‰ and +0.3 ± 0.4‰, respectively. The CO₂ released by root respiration was ¹³C depleted by about −2.1 ± 2.2‰ for C₃ plants and −1.3 ± 2.4‰ for C₄ plants compared to root tissue. However, only a very few studies investigated ¹³C fractionation by root respiration. This urgently calls for further research. In soils developed under C₃ vegetation, the microbial biomass was ¹³C enriched by +1.2 ± 2.6‰ and microbial CO₂ was also ¹³C enriched by +0.7 ± 2.8‰ compared to SOM. This discrimination pattern suggests preferential utilization of ¹³C-enriched substances by microorganisms, but a respiration of lighter compounds from this fraction. The δ¹³C signature of the microbial pool is composed of metabolically active and dormant microorganisms; the respired CO₂, however, derives mainly from active organisms. This discrepancy and the preferential substrate utilization explain the δ¹³C differences between microorganisms and CO₂ by an ‘apparent’ ¹³C discrimination. Preferential consumption of easily decomposable substrates and less negative δ¹³C values were common for substances with low C/N ratios. Preferential substrate utilization was more important for C₃ soils because, in C₄ soils, microbial respiration strictly followed kinetics, i.e. microorganisms incorporated heavier C (? = +1.1‰) and respired lighter C (? = −1.1‰) than SOM. Temperature and precipitation had no significant effect on the ¹³C fractionation in these processes in C₃ soils. Increasing temperature and decreasing precipitation led, however, to increasing δ¹³C of soil C pools.Based on these ¹³C fractionations we developed a number of consequences for C partitioning studies using ¹³C natural abundance. In the framework of standard isotope mixing models, we calculated CO₂ partitioning using the natural-¹³C-labelling approach at a vegetation change from C₃ to C₄ plants assuming a root-derived fraction between 0% and 100% to total soil CO₂. Disregarding any ¹³C fractionation processes, the calculated results deviated by up to 10% from the assumed fractions. Accounting for ¹³C fractionation in the standard deviations of the C₄ source and the mixing pool did not improve the exactness of the partitioning results; rather, it doubled the standard errors of the CO₂ pools. Including ¹³C fractionations directly into the mass balance equations reproduced the assumed CO₂ partitioning exactly. At the end, we therefore give recommendations on how to consider ¹³C fractionations in research on carbon flows between plants, microorganisms, and soil.     

CO2 efflux by rapid decomposition of low molecular organic substances in soils

Decomposition rates of the [2-¹⁴C]-glucose and [2-¹⁴C]-glycine in four different soils of the long-term field trial of Moscow were investigated in a 3-months laboratory experiment in which ¹⁴CO₂ respiration was measured. A model with three decomposition components and two distribution parameters was developed and validated with the data of the experiment. The decay rate constants of free [2-¹⁴C]-glucose (4–32 day^-1) were slower than those of [2-¹⁴C]-glycine (16–44 day^-1). The calculated use efficiency for microbial biosynthesis of the second carbon atom was 47% for glucose and 31% for glycine. The potential half-life of labelled carbon in the microbial soil biomass ranged from 0.6 to 4.4 days, depending on the soil type and the initial amount of added substrate. The calculated total utilisation of carbon by the soil biomass from glycine was about 2–5 times lower than that of glucose.The modelled ¹⁴C incorporation into the microbial soil biomass reached its maximum on the first day of the incubation experiment and did not exceed 22% of the ¹⁴C input. Both of the investigated substances decomposed most rapidly in the soil samples from sites that have not being fertilised with organic or mineral fertilisers during an 81-years period.