外源氮添加对土壤微生物残体积累动态的影响－基于Meta分析

微生物残体是土壤有机碳库的重要贡献者。为明确外源氮添加对土壤微生物残体积累动态的影响,本文收集整理了1980—2020年已发表的文献,共选取122组试验观测数据,利用整合分析方法（Meta-analysis）,以微生物残体标识物－氨基糖为目标组分,定量分析了不同种类和数量的外源氮添加对土壤中微生物来源细胞残体积累数量和组成比例的影响,并系统解析其主要影响因素。结果表明:外源氮添加（0 ~ 6000 kg hm?1）对微生物细胞残体的积累有显著的促进作用,并能引起土壤中真菌和细菌来源细胞残体相对比例发生明显变化。与不加氮对照相比,氮添加使土壤氨基糖总量增加27%,其中氨基葡萄糖、氨基半乳糖和胞壁酸含量分别增加22.5%、29.8%和19.0%。同时,不同种类外源氮素添加对氨基糖积累特征的影响也有所不同,表现为有机氮（如动物厩肥）比无机氮添加对氨基糖积累的促进作用更大。此外,氮添加对氨基糖的影响程度还与土壤自身的碳氮比、土地利用类型和自然降雨量等环境因子密切相关。其中是否添加碳源对微生物残体的响应有较大影响,表现为:无碳源添加会降低土壤氨基糖葡萄糖和胞壁酸对氮添加的响应,削弱了微生物残体对土壤有机质的贡献比例;而氮源同时配合碳源添加条件下,土壤氨基糖积累量显著高于单一氮源添加的处理,说明氮添加对微生物残体积累的影响存在着碳氮耦合效应。     

Taxon-specific responses of soil bacteria to the addition of low level C inputs

The addition of small or trace amounts of carbon to soils can result in the release of 2-5 times more C as CO₂ than was added in the original solution. The identity of the microorganisms responsible for these so-called trigger effects remains largely unknown. This paper reports on the response of individual bacterial taxa to the addition of a range of ¹⁴C-glucose concentrations (150, 50 and 15 and 0 μg C g⁻¹ soil) similar to the low levels of labile C found in soil. Taxon-specific responses were identified using a modification of the stable isotope probing (SIP) protocol and the recovery of [¹⁴C] labelled ribosomal RNA using equilibrium density gradient centrifugation. This provided good resolution of the ‘heavy’ fractions ([¹⁴C] labelled RNA) from the ‘light’ fractions ([¹²C] unlabelled RNA). The extent of the separation was verified using autoradiography. The addition of [¹⁴C] glucose at all concentrations was characterised by changes in the relative intensity of particular bands. Canonical correspondence analysis (CCA) showed that the rRNA response in both the ‘heavy’ and ‘light’ fractions differed according to the concentration of glucose added but was most pronounced in soils amended with 150 μg C g⁻¹ soil. In the ‘heavy RNA’ fractions there was a clear separation between soils amended with 150 μg C g⁻¹ soil and those receiving 50 and 15 μg C g⁻¹ soil indicating that at low C inputs the microbial community response is quite distinct from that seen at higher concentrations. To investigate these differences further, bands that changed in relative intensity following amendment were excised from the DGGE gels, reamplified and sequenced. Sequence analysis identified 8 taxa that responded to glucose amendment (Bacillus, Pseudomonas, Burkholderia, Bradyrhizobium, Actinobacteria, Nitrosomonas, Acidobacteria and an uncultured β-proteobacteria). These results show that radioisotope probing (RNA-RIP) can be used successfully to study the fate of labile C substrates, such as glucose, in soil.     

Effect of soil copper on the response of soil fungal communities to the addition of plant residues

Organic residues provide the fundamental energy supply supporting soil fungal communities. Provision of adequate energy is required for soil microbial communities to adapt and function in the presence of ecological stress, such as copper (Cu) contamination. However, contamination can also lead to decreased ecological fitness of microorganisms, limiting their ability to access substrates. Thus, complex interactions exist between substrates, metals, energy supply/accessibility, fungal communities and their processes, and these have implications for ecosystem processes. We investigated the interaction between energy resources and Cu tolerance on soil fungal communities, including Fusarium and Trichoderma (model disease causing and beneficial genera). Using quantitative PCR and DGGE fingerprinting, the effects of increasing soil Cu levels (0 to >3000 mg Cu kg⁻¹ soil) on size and structure of soil fungal communities were tested under basal and plant-residue (medic; Medicago trunculata) added conditions. The interaction between increasing soil Cu levels and the addition of plant resources on fungal community structure was tested using multivariate analysis. The relative size (DNA copies per unit of soil DNA) of soil fungal communities, including Trichoderma and Fusarium, significantly (P < 0.05) increased (94% and 32% respectively) with addition of medic to soil. In medic-applied samples, the bacterial to fungal ratio decreased, demonstrating the selective influence of the cellulose-rich substrate on the fungal community. Under the high nutrient conditions fungal DNA increased as a fraction of the total soil DNA, demonstrating the tolerance of fungi to Cu (relative to other microbiota) given adequate energy resources. Copper had no impact on the abundance of Fusarium or Trichoderma, but significantly affected community structure (PERMANOVA; P < 0.05). With increasing Cu, species selection and replacement could be observed, particularly in soils where medic had been included. Plant residue addition itself was a highly selective factor affecting the structure of communities of Trichoderma and Fusarium (P < 0.05). The effects of increasing Cu could be seen in both medic and basal soils for Trichoderma, but only in the basal treatments for soil Fusarium. This was due to very low dispersion in Fusarium community structure in the medic-added treatment (PERMDISP; P < 0.05). The results show the interactive influence of organic matter inputs and heavy metal contamination on size and structure of soil fungal communities. The data show that species selection and replacement is an important mechanism for community adaptation to increasing levels of soil Cu, and this mechanism can be influenced by addition of resources to the soil.     

Titanium oxide nanoparticle effects on composition of soil microbial communities and plant performance

We examined the effect of TiO₂ nanoparticles (NPs) on the growth of maize and soybean plants and associated soil microbial communities. Plants were grown in a greenhouse, and low levels of undoped or nitrogen-doped TiO₂ NPs were applied. Plant growth and nutrient content were determined, and effects of NPs on composition of soil microbial communities were examined using terminal restriction fragment length polymorphism analysis (TRFLP) of rDNA. We found no significant effects of TiO₂ NPs on plant growth, nutrient content, or the composition of bacterial communities within the rhizosphere. However, arbuscular mycorrhizal fungal communities were affected by application of undoped and nitrogen-doped TiO₂ NPs. This observation may be partially attributed to the small but significant TiO₂ NP uptake levels in the root tissues of both plants. Our results suggest that even low concentrations of TiO₂ NPs may influence some important groups of soil microbes, such as mycorrhizal fungi, but changes in the composition of microbial communities may not affect plant growth under conditions of adequate moisture and nutrients.     

Effects of glyphosate on rhizosphere soil microbial communities under two different plant compositions by cultivation-dependent and -independent methodologies

We studied, under two different plant compositions, the short-term effects of glyphosate on rhizosphere soil microbial communities through the utilization of cultivation-dependent and -independent techniques. A short-term pot study was carried out using factorial treatments that included two different compositions of forage plant species (triticale versus a mixture of triticale and pea) and two concentrations of glyphosate (50 and 500 mg active ingredient kg⁻¹ soil, as a commercial formulation, Roundup Plus) arranged in a completely randomized design experiment with four replicates. Control plants (no glyphosate added) were clipped in an attempt to compare two methods of weed control (manual = clipping; chemical = herbicide treatment). Rhizosphere soil was sampled 15 and 30 days after glyphosate treatment and the following soil components were determined: potentially mineralizable nitrogen, ammonium content, community-level physiological profiles using Biolog Ecoplates™, DNA microbial biomass and genotype diversity by means of PCR-DGGE. Fifteen days after herbicide treatment, a glyphosate-induced stimulation of the activity and functional diversity of the cultivable portion of the heterotrophic soil microbial community was observed, most likely due to glyphosate acting as an available source of C, N and P. On the other hand, 30 days after herbicide treatment, both the activity and diversity of the rhizosphere soil microbial communities showed an inconsistent response to glyphosate addition. Apart from its intended effect on plants, glyphosate had non-target effects on the rhizosphere soil microbial community which were, interestingly, more enhanced in triticale than in “triticale + pea” pots. Biolog™ was more sensitive than PCR-DGGE to detect changes in soil microbial communities induced by glyphosate and plant composition.     

Precipitation modifies the effects of warming and nitrogen addition on soil microbial communities in northern Chinese grasslands

Terrestrial ecosystems experience simultaneous shifts in multiple drivers of global change, which can interactively affect various resources. The concept that different resources co-limit plant productivity has been well studied. However, co-limitation of soil microbial communities by multiple resources has not been as thoroughly investigated. Specifically, it is not clearly understood how microbial communities respond to shifts in multiple interacting resources such as water, temperature, and nitrogen (N), in the context of global change. To test the effects of these various resources on soil microorganisms, we established a field experiment with temperature and N manipulation in three grasslands of northern China, where there is a decrease in precipitation from east to west across the region. We found that microbial responses to temperature depended upon seasonal water regimes in these temperate steppes. When there was sufficient water present, warming had positive effects on soil microorganisms, suggesting an interaction between water and increases in temperature enhanced local microbial communities. When drought or alternating wet–dry stress occurred, warming had detrimental effects on soil microbial communities. Our results also provide clear evidence for serial co-limitation of microorganisms by water and N at the functional group and community levels, where water is a primary limiting factor and N addition positively affects soil microorganisms only when water is sufficient. We predict that future microbial responses to changes in temperature and N availability could be seasonal or exist only in non-drought years, and will strongly rely on future precipitation regimes.     

Functional and molecular responses of soil microbial communities under differing soil management practices

The effects of soil management on some microbiological properties and soil bacterial community structure were evaluated. Two field sites with the same soil type, located on the same geographic area adjacent to one other, have received different soil management practices and cultivation. One site has been subjected for 20 years to intensive horticulture under conventional tillage and irrigation with low quality salt-rich water; the second field site has been uncultivated for a long period and was turned to organic farming practices over the last 5 years and is currently cultivated with fruit orchard. Total bacterial counts, microbial ATP, microbial community metabolic (BIOLOG^®) profiles, and DNA fingerprinting by PCR-DGGE were determined. Two-way ANOVA revealed that total bacterial counts were not significantly (P>0.3) affected by the two different management practices; ATP content was consistently and significantly (P<0.001) lower in salt-water irrigated soil than in organic soil at the three sampling times. The cluster analysis of community level physiological profiles indicated that microbial communities were much more uniform in organic soil than in irrigated one, suggesting that salt-water irrigation could have affected the size of the microbial population, its metabolic activities, as well as its composition. Molecular patterns fitted the BIOLOG^® profile diversity. In particular, at any sampling time, PCR-DGGE patterns of bacterial DNA, extracted by an indirect method, significantly discriminated irrigated from organic soil samples. The PCR-DGGE patterns of total soil DNA, extracted by a direct method, showed a moderate to significant variation among irrigated and organic soil samples. Biochemical, microbiological and molecular data contributed to evidence a significantly different response of indigenous microflora to soil management by using saline water or organic farming.     

不同筛分方式对土壤团聚体及土壤微生物群落的影响

探究土壤团聚体组成、稳定性、有机碳含量及微生物数量在不同土壤筛分方式下的变化,对团聚体筛分方式进行优化。以黑土为研究对象,设置不同筛分方式处理,比较干土干筛、干土湿筛、润土干筛、润土湿筛4种筛分方式的优缺点。结果表明,润土干筛比干土湿筛处理≥2 mm粒径的土壤团聚体高出约85.97%,而在2～0.25、≤0.25 mm粒级中团聚体所占比例最高的干土湿筛比最低的润土干筛分别高出约80.90%、91.82%。>0.25 mm水稳性团聚体含量(R_0.25)在润土干筛下达到最高,为99.35%,该指标与土壤团聚体稳定性成正相关,其值越高则表示土壤抗蚀能力越强。干土湿筛处理下各粒级团聚体土壤有机碳分布平均。干土湿筛处理下,土壤细菌和放线菌的数量最高。而润土干筛处理下土壤真菌数量较高,与润土湿筛、干土干筛、干土湿筛3个处理方式相比有显著性差异。综上发现,干土湿筛处理下土壤团聚体稳定性更好,土壤微生物数量较高,在一定程度上对土壤结构、土壤质量起到优化的作用。     

Dissimilar response of plant and soil biota communities to long-term nutrient addition in grasslands

The long-term effect of fertilizers on plant diversity and productivity is well known, but long-term effects on soil biota communities have received relatively little attention. Here, we used an exceptional long-lasting (>40 years) grassland fertilization experiment to investigate the long-term effect of Ca, N, PK, and NPK addition on the productivity and diversity of both vegetation and soil biota. Whereas plant diversity increased by liming and decreased by N and NPK, the diversity of nematodes, collembolans, mites, and enchytraeids increased by N, PK, or NPK. Fertilization with NPK and PK increased plant biomass and biomass of enchytraeids and collembolans. Biomass of nematodes and earthworms increased by liming. Our results suggest that soil diversity might be driven by plant productivity rather than by plant diversity. This may imply that the selection of measures for restoring or conserving plant diversity may decrease soil biota diversity. This needs to be tested in future experiments.     

Response of microbial communities to different doses of chromate in soil microcosms

The toxic effect of chromate on soil microbial communities is not well documented, although microorganisms control biogeochemical cycling, contribute to formation of soil structure, regulate the fate of organic matter applied to soil. In this study the effects of short- and middle-term chromate on the soil microbial community were investigated. The shifts in the size and in the diversity of culturable heterotrophic bacterial community, the resistance to Cr(VI) of heterotrophic bacteria, the presence of cyanobacteria, the activity of 19 enzymes, and the ATP content were monitored over time (120 days) in soil microcosms artificially contaminated with three concentrations of chromate (50, 250 and 1000 mg kg⁻¹ soil). The chromate contamination affected the structure and the diversity of the soil bacterial community. Bacterial strains isolated from the microcosm contaminated with the highest concentration of chromate were identified by 16S rDNA gene sequencing. All isolates belonged to the genus Pseudomonas, were able to reduce Cr(VI), and showed a high resistance to chromate. To our knowledge, this is the first report that shows Pseudomonas strains having the capability to resist up to 40 mM of Cr(VI) on minimal medium. The cyanobacterial group was more sensitive to chromate contamination than culturable heterotrophic bacteria. No cyanobacterial growth was detected in enrichment cultures from the soil polluted with the highest chromate concentration. Some enzymes were inhibited by high concentrations of chromate, whereas others were stimulated. The ATP content in microcosms was strongly affected by chromate. We conclude that the soil microbial community responds to chromate pollution through changes in community structure, in metabolic activity, and in selection for Cr(VI)-resistance.     

Rapid effects of plant species diversity and identity on soil microbial communities in experimental grassland ecosystems

Changes in plant community structure, including the loss of plant diversity may affect soil microbial communities. To test this hypothesis, plant diversity and composition were experimentally varied in grassland plots cultivated with monocultures or mixtures of 2, 3 or 4 species. We tested the effects of monocultures versus mixtures and of plant species composition on culturable soil bacterial activity, number of substrates used and catabolic diversity, microbial biomass N, microbial respiration, and root biomass. These properties were all measured 10 months after seeding the experiment. Soil bacterial activity, number of substrates used and catabolic diversity were measured in the different plant communities using BIOLOG GN and GP microplates, which are redox-based tests measuring capacity of soil culturable bacteria to use a variety of organic substrates. Microbial biomass N, microbial respiration, and root biomass were insensitive to plant diversity. Culturable soil microbial activity, substrates used and diversity declined with declining plant diversity. Their activity, number of substrates used and diversity were significantly higher in plots with 3 and 4 plant species than in monocultures and in plots with 2 species. There was also an effect of plant species composition. Culturable soil microbial activity and diversity was higher in the four-species plant community than in any of the plant monocultures suggesting that the effect of plant diversity could not be explained by the presence of a particular plant species. Our results showed that changes in plant diversity and composition in grassland ecosystems lead to a rapid response of bacterial activity and diversity.     

Differential responses of total and active soil microbial communities to long-term experimental N deposition

The relationship between total and metabolically active soil microbial communities can provide insight into how these communities are impacted by environmental change, which may impact the flow of energy and cycling of nutrients in the future. For example, the anthropogenic release of biologically available N has dramatically increased over the last 150 years, which can alter the processes controlling C storage in terrestrial ecosystems. In a northern hardwood forest ecosystem located in Michigan, USA, nearly 20 years of experimentally increased atmospheric N deposition has reduced forest floor decay and increased soil C storage. A microbial mechanism underlies this response, as compositional changes in the soil microbial community have been concomitantly documented with these biogeochemical changes. Here, we co-extracted DNA and RNA from decaying leaf litter to determine if experimental atmospheric N deposition has lowered the diversity and altered the composition of the whole communities of bacteria and fungi (i.e., DNA-based) and well as its active members (i.e., RNA-based). In our experiment, experimental N deposition did not affect the composition, diversity, or richness of the total forest floor fungal community, but did lower the diversity (−8%), as well as altered the composition of the active fungal community. In contrast, neither the total nor active forest floor bacterial community was significantly affected by experimental N deposition. Our results suggest that future rates of atmospheric N deposition can fundamentally alter the organization of the saprotrophic soil fungal community, key mediators of C cycling in terrestrial environments.     

Negative priming of native soil organic carbon mineralization by oilseed biochars of contrasting quality

Oilseed‐derived biochar, a by‐product of pyrolysis for biodiesel production, is richer in aliphatic compounds than the commonly studied wood‐derived biochar, affecting both its mineralization in soil and its interaction with native soil organic carbon (nSOC). Here, we investigated the soil C sequestration potential of three different oilseed biochars derived from C₃ plant material: soyabean, castor bean and jatropha cake. The chemical composition of these biochars was determined by elemental analysis (CHN) and ¹³C NMR spectroscopy. The cumulative CO₂ efflux from 30‐day laboratory incubations of biochar mixed with a sandy soil containing nSOC from C₄ plants was measured as a proxy for mineralization rate. The relative contribution of each source to CO₂ production was calculated based on the ¹³C‐signatures of total CO₂ efflux and the source materials (soil and biochars). Our results showed that: (i) castor bean biochar contained relatively large amounts of aliphatic compounds, resulting in a greater mineralization rate than soyabean and jatropha biochars; (ii) CO₂ efflux from the soil‐biochar mixtures originated mostly from the biochars, suggesting that these biochars contain rapidly decomposable compounds; and (iii) all three oilseed biochars decelerated nSOC mineralization. This negative priming effect appeared to be caused by different factors. We conclude that oilseed biochars have the potential to increase soil C stocks directly and increase soil C sequestration indirectly in the short term through negative priming of nSOC mineralization.     

Mineralization and microbial biomass formation in upland soil amended with some tropical plant residues at different temperatures

A model experiment was carried out at 15, 25, and 35°C to investigate the changes in microbial biomass and the pattern of mineralization in upland soil during 8 weeks following the addition of 8 organic materials including 6 tropical plant residues, ipil ipil (Leucaena leucocephala), azolla (Azolla pinnata), water hyacinth (Eichhornia crassipes), dhaincha (Sesbania rostrata), cowpea (Vigna unguiculata), and sunhemp (Crotalaria juncea). The amounts of CO₂-C evolved and inorganic N produced at 35°C were about 2 times larger than those at 15°C. At any temperature, the flush decomposition of C was observed within the first week and thereafter the rate of mineralization became relatively slow. A negative correlation was observed between inorganic N and C/N ratios of the added organic materials. The relationships between the amounts of cellulose or cellulose plus hemicellulose and the amount of mineralized N of the added organic materials were also negative.

The changes in the microbial biomass were affected by temperatures. The amount of biomass C and N was maximum after 42 d of incubation at 15°C, and after 7 d at 25 and 35°C, and thereafter decreased. The rate of biomass decline was slower at 15°C and faster at 35°C than at 25°C. Regardless of the temperatures, the addition of organic materials enhanced microbial biomass formation throughout the incubation periods.     

Comparative analysis of soil microbial communities and their responses to the short-term drought in bog,fen, and riparian wetlands

The frequency of drought is anticipated to increase in wetland ecosystems as global warming intensifies. However, information on microbial communities involved in greenhouse gas emissions and their responses to drought remains sparse. We compared the gene abundance of eubacterial 16S rRNA, nitrite reductase (nirS) and methyl coenzyme M reductase (mcrA), and the diversity and composition of eubacteria, methanogens and denitrifiers among bog, fen and riparian wetlands. The gene abundance, diversity and composition significantly differed among wetlands (p < 0.01) with the exception of the diversity of methanogens. The gene abundance was ranked in the order of the bog = fen > riparian wetland, whereas the diversity was in the riparian wetland ≥ fen > bog. In addition, we conducted a short-term drought experiment and compared microbial communities between control (water-logged) and drought (?15 cm) treatments. Drought led to significant decline in the gene abundance in the bog (16S rRNA, nirS, mcrA) (p < 0.01) and fen (16S rRNA, nirS) (p < 0.05), but not in the riparian wetland. There were no differences in the diversity and composition of denitrifiers and methanogens at all sites following drought. Our results imply that denitrifiers and methanogens inhabiting bogs and fens would suffer from short-term droughts, but remain unchanged in riparian wetlands.     

重复添加植物残体后土壤微生物活性和生物量对盐度的响应特征

Microbial adaptation to salinity can be achieved through synthesis of organic osmolytes,which requires high amounts of energy;however,a single addition of plant residues can only temporarily improve energy supply to soil microbes.Therefore,a laboratory incubation experiment was conducted to evaluate the responses of soil microbes to increasing salinity with repeated additions of plant residues using a loamy sand soil with an electrical conductivity in saturated paste extract（EC_e） of 0.6 dS m^-1.The soil was kept non-saline or salinized by adding different amounts of NaCl to achieve EC_e of 12.5,25.0 and 50.0 dS m^-1.The non-saline soil and the saline soils were amended with finely ground pea residues at two rates equivalent to 3.9 and 7.8 g C kg^-1 soil on days 0,15 and29.The soils receiving no residues were included as a control.Cumulative respiration per g C added over 2 weeks after each residue addition was always greater at 3.9 than 7.8 g C kg^-1 soil and higher in the non-saline soil than in the saline soils.In the saline soils,the cumulative respiration per g C added was higher after the second and third additions than after the first addition except with3.9 g C kg^-1 at EC_e of 50 dS m^₁.Though with the same amount of C added(7.8 g C kg^-1),salinity reduced soil respiration to a lesser extent when 3.9 g C kg^-1 was added twice compared to a single addition of 7.8 g C kg^-1.After the third residue addition,the microbial biomass C concentration was significantly lower in the soils with EC_e of 25 and 50 dS m^₁ than in the non-saline soil at3.9 g C kg^-1,but only in the soil with EC_e of 50 dS m^-1 at 7.8 g C kg^-1.We concluded that repeated residue additions increased the adaptation of soil microbial community to salinity,which was likely due to high C availability providing microbes with the energy needed for synthesis of organic osmolytes.