Sources of CO2 efflux from soil and review of partitioning methods

Five main biogenic sources of CO₂ efflux from soils have been distinguished and described according to their turnover rates and the mean residence time of carbon. They are root respiration, rhizomicrobial respiration, decomposition of plant residues, the priming effect induced by root exudation or by addition of plant residues, and basal respiration by microbial decomposition of soil organic matter (SOM). These sources can be grouped in several combinations to summarize CO₂ efflux from the soil including: root-derived CO₂, plant-derived CO₂, SOM-derived CO₂, rhizosphere respiration, heterotrophic microbial respiration (respiration by heterotrophs), and respiration by autotrophs. These distinctions are important because without separation of SOM-derived CO₂ from plant-derived CO₂, measurements of total soil respiration have very limited value for evaluation of the soil as a source or sink of atmospheric CO₂ and for interpreting the sources of CO₂ and the fate of carbon within soils and ecosystems. Additionally, the processes linked to the five sources of CO₂ efflux from soil have various responses to environmental variables and consequently to global warming. This review describes the basic principles and assumptions of the following methods which allow SOM-derived and root-derived CO₂ efflux to be separated under laboratory and field conditions: root exclusion techniques, shading and clipping, tree girdling, regression, component integration, excised roots and insitu root respiration; continuous and pulse labeling, ¹³C natural abundance and FACE, and radiocarbon dating and bomb-¹⁴C. A short sections cover the separation of the respiration of autotrophs and that of heterotrophs, i.e. the separation of actual root respiration from microbial respiration, as well as methods allowing the amount of CO₂ evolved by decomposition of plant residues and by priming effects to be estimated. All these methods have been evaluated according to their inherent disturbance of the ecosystem and C fluxes, and their versatility under various conditions. The shortfalls of existing approaches and the need for further development and standardization of methods are highlighted.     

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.     

Rhizodeposition: Its contribution to microbial growth and carbon and nitrogen turnover within the rhizosphere

A greenhouse rhizobox experiment was carried out to investigate the fate and turnover of ¹³C‐ and ¹⁵N‐labeled rhizodeposits within a rhizosphere gradient from 0–8 mm distance to the roots of wheat. Rhizosphere soil layers from 0–1, 1–2, 2–3, 3–4, 4–6, and 6–8 mm distance to separated roots were investigated in an incubation experiment (42 d, 15°C) for changes in total C and N and that derived from rhizodeposition in total soil, in soil microbial biomass, and in the 0.05 M K₂SO₄–extractable soil fraction. CO₂‐C respiration in total and that derived from rhizodeposition were measured from the incubated rhizosphere soil samples. Rhizodeposition C was detected in rhizosphere soil up to 4–6 mm distance from the separated roots. Rhizodeposition N was only detected in the rhizosphere soils up to 3–4 mm distance from the roots. Microbial biomass C and N was increased with increasing proximity to the separated roots. Beside ¹³C and ¹⁵N derived from rhizodeposits, unlabeled soil C and N (native SOM) were incorporated into the growing microbial biomass towards the roots, indicating a distinct acceleration of soil organic matter (SOM) decomposition and N immobilization into the growing microbial biomass, even under the competition of plant growth. During the soil incubation, microbial biomass C and N decreased in all samples. Any decrease in microbial biomass C and N in the incubated rhizosphere soil layers is attributed mainly to a decrease of unlabeled (native) C and N, whereas the main portion of previously incorporated rhizodeposition C and N during the plant growth period remained immobilized in the microbial biomass during the incubation. Mineralization of native SOM C and N was enhanced within the entire investigated rhizosphere gradient. The results indicate complex interactions between substrate input derived from rhizodeposition, microbial growth, and accelerated C and N turnover, including the decomposition of native SOM (i.e., rhizosphere priming effects) at a high spatial resolution from the roots.     

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.     

Plant biomass influences rhizosphere priming effects on soil organic matter decomposition in two differently managed soils

We used a continuous labeling method of naturally ¹³C-depleted CO₂ in a growth chamber to test for rhizosphere effects on soil organic matter (SOM) decomposition. Two C3 plant species, soybean (Glycine max) and sunflower (Helianthus annus), were grown in two previously differently managed soils, an organically farmed soil and a soil from an annual grassland. We maintained a constant atmospheric CO₂ concentration at 400±5 ppm and δ¹³C signature at −24.4‰ by regulating the flow of naturally ¹³C-depleted CO₂ and CO₂-free air into the growth chamber, which allowed us to separate new plant-derived CO₂-C from original soil-derived CO₂-C in soil respiration. Rhizosphere priming effects on SOM decomposition, i.e., differences in soil-derived CO₂-C between planted and non-planted treatments, were significantly different between the two soils, but not between the two plant species. Soil-derived CO₂-C efflux in the organically farmed soil increased up to 61% compared to the no-plant control, while the annual grassland soil showed a negligible increase (up to 5% increase), despite an overall larger efflux of soil-derived CO₂-C and total soil C content. Differences in rhizosphere priming effects on SOM decomposition between the two soils could be largely explained by differences in plant biomass, and in particular leaf biomass, explaining 49% and 74% of the variation in primed soil C among soils and plant species, respectively. Nitrogen uptake rates by soybean and sunflower was relatively high compared to soil C respiration and associated N mineralization, while inorganic N pools were significantly depleted in the organic farm soil by the end of the experiment. Despite relatively large increases in SOM decomposition caused by rhizosphere effects in the organic farm soil, the fast-growing soybean and sunflower plants gained little extra N from the increase in SOM decomposition caused by rhizosphere effects. We conclude that rhizosphere priming effects of annual plants on SOM decomposition are largely driven by plant biomass, especially in soils of high fertility that can sustain high plant productivity.     

Moisture modulates rhizosphere effects on C decomposition in two different soil types

While it is well known that soil moisture directly affects microbial activity and soil organic matter (SOM) decomposition, it is unclear if the presence of plants alters these effects through rhizosphere processes. We studied soil moisture effects on SOM decomposition with and without sunflower and soybean. Plants were grown in two different soil types with soil moisture contents of 45% and 85% of field capacity in a greenhouse experiment. We continuously labeled plants with depleted ¹³C, which allowed us to separate plant-derived CO₂-C from original soil-derived CO₂-C in soil respiration measurements. We observed an overall increase in soil-derived CO₂-C efflux in the presence of plants (priming effect) in both soils. On average a greater priming effect was found in the high soil moisture treatment (up to 76% increase in soil-derived CO₂-C compared to control) than in the low soil moisture treatment (up to 52% increase). Greater plant-derived CO₂-C and plant biomass in the high soil moisture treatment contributed to greater priming effects, but priming effects remained significantly higher in the high moisture treatment than in the low moisture treatment after correcting for the effects of plant-derived CO₂-C and plant biomass. The response to soil moisture particularly occurred in the sandy loam soil by the end of the experiment. Possibly, production of root exudates increased with increased soil moisture content. Root exudation of labile C may also have become more effective in stimulating microbial decomposition in the higher soil moisture treatment and sandy loam soil. Our results indicate that moisture conditions significantly modulate rhizosphere effects on SOM decomposition.     

Root and rhizomicrobial respiration: A review of approaches to estimate respiration by autotrophic and heterotrophic organisms in soil

Partitioning the root‐derived CO₂ efflux from soil (frequently termed rhizosphere respiration) into actual root respiration (RR, respiration by autotrophs) and rhizomicrobial respiration (RMR, respiration by heterotrophs) is crucial in determining the carbon (C) and energy balance of plants and soils. It is also essential in quantifying C sources for rhizosphere microorganisms and in estimation of the C contributing to turnover of soil organic matter (SOM), as well as in linking net ecosystem production (NEP) and net ecosystem exchange (NEE). Artificial‐environment studies such as hydroponics or sterile soils yield unrealistic C‐partitioning values and are unsuitable for predicting C flows under natural conditions. To date, several methods have been suggested to separate RR and RMR in nonsterile soils: 1) component integration, 2) substrate‐induced respiration, 3) respiration by excised roots, 4) comparison of root‐derived ¹⁴CO₂ with rhizomicrobial ¹⁴CO₂ after continuous labeling, 5) isotope dilution, 6) model‐rhizodeposition technique, 7) modeling of ¹⁴CO₂ efflux dynamics, 8) exudate elution, and 9) δ¹³C of CO₂ and microbial biomass. This review describes the basic principles and assumptions of these methods and compares the results obtained in the original papers and in studies designed to compare the methods. The component‐integration method leads to strong disturbance and non‐proportional increase of CO₂ efflux from different sources. Four of the methods (5 to 8) are based on the pulse labeling of shoots in a ¹⁴CO₂ atmosphere and subsequent monitoring of ¹⁴CO₂ efflux from the soil. The model‐rhizodeposition technique and exudate‐elution procedure strongly overestimate RR and underestimate RMR. Despite alternative assumptions, isotope dilution and modeling of ¹⁴CO₂‐efflux dynamics yield similar results. In crops and grasses (wheat, ryegrass, barley, buckwheat, maize, meadow fescue, prairie grasses), RR amounts on average to 48±5% and RMR to 52±5% of root‐derived CO₂. The method based on the ¹³C isotopic signature of CO₂ and microbial biomass is the most promising approach, especially when the plants are continuously labeled in ¹³CO₂ or ¹⁴CO₂ atmosphere. The “difference” methods, i.e., trenching, tree girdling, root‐exclusion techniques, etc., are not suitable for separating the respiration by autotrophic and heterotrophic organisms because the difference methods neglect the importance of microbial respiration of rhizodeposits.     

Microbial community abundance and structure are determinants of soil organic matter mineralisation in the presence of labile carbon

Altered rates of native soil organic matter (SOM) mineralisation in the presence of labile C substrate (‘priming’), is increasingly recognised as central to the coupling of plant and soil-biological productivity and potentially as a key process mediating the C-balance of soils. However, the mechanisms and controls of SOM-priming are not well understood. In this study we manipulated microbial biomass size and composition (chloroform fumigation) and mineral nutrient availability to investigate controls of SOM-priming. Effects of applied substrate (¹³C-glucose) on mineralisation of native SOM were quantified by isotopic partitioning of soil respiration. In addition, the respective contributions of SOM-C and substrate-derived C to microbial biomass carbon (MBC) were quantified to account for pool-substitution effects (‘apparent priming’). Phospholipid fatty acid (PLFA) profiles of the soils were determined to establish treatment effects on microbial community structure, while the ¹³C-enrichment of PLFA biomarkers was used to establish pathways of substrate-derived C-flux through the microbial communities. The results indicated that glucose additions increased SOM-mineralisation in all treatments (positive priming). The magnitude of priming was reduced in fumigated soils, concurrent with reduced substrate-derived C-flux through putative SOM-mineralising organisms (fungi and actinomycetes). Nutrient additions reduced the magnitude of positive priming in non-fumigated soils, but did not affect the distribution of substrate-derived C in microbial communities. The results support the view that microbial community composition is a determinant of SOM-mineralisation, with evidence that utilisation of labile substrate by fungal and actinomycete (but not Gram-negative) populations promotes positive SOM-priming.     

Soil physicochemical and microbial drivers of temperature sensitivity of soil organic matter decomposition under boreal forests

Soil organic matter(SOM)in boreal forests is an important carbon sink.The aim of this study was to assess and to detect factors controlling the temperature sensitivity of SOM decomposition.Soils were collected from Scots pine,Norway spruce,silver birch,and mixed forests(O horizon)in northern Finland,and their basal respiration rates at five different temperatures(from 4 to 28℃)were measured.The Q_(10) values,showing the respiration rate changes with a 10℃ increase,were calculated using a Gaussian function and were based on temperature-dependent changes.Several soil physicochemical parameters were measured,and the functional diversity of the soil microbial communities was assessed using the MicroResp?method.The temperature sensitivity of SOM decomposition differed under the studied forest stands.Pine forests had the highest temperature sensitivity for SOM decomposition at the low temperature range(0–12℃).Within this temperature range,the Q_(10) values were positively correlated with the microbial functional diversity index(H'_(mic))and the soil C-to-P ratio.This suggested that the metabolic abilities of the soil microbial communities and the soil nutrient content were important controls of temperature sensitivity in taiga soils.     

Effect of plant roots on carbon metabolism of soil microbial biomass

The utilization of plant- and soil-C by the microbial biomass in the rhizosphere of maize plants was investigated as a function of root proximity. The plants were cultivated in pots with divided root chambers and their shoots supplied with ¹⁴CO₂ for 23 days. Subsequently the individual soil zones were analyzed for organic C, ¹⁴C, biomass C and biomass ¹⁴C. Plant roots induced a 197% increase in microbial biomass and a 5.4% decrease in soil organic C compared with an 1.2% decrease in the unplanted control soil. The contributions of plant- and soil-C to this increased microbial growth amounted to 68% and 32% respectively. Biomass-¹⁴C corresponded to 1.6% of the total photosynthetically fixed ¹⁴C, to about 15% of the organic ¹⁴C-input into the rhizosphere and to 58% of the plant carbon remaining in soil after the removal of roots. 20% of this biomass-¹⁴C was found outside the immediate root zone. These results demonstrate that growing roots are a significant C-source for the microbial biomass and render an additional fraction of soil-C available to microbial utilization. The efficiency of C-utilization by the rhizosphere biomass is lower than values obtained with liquid cultures in laboratory experiments. The supply of plant-C to the microbial biomass outside the immediate root vicinity indicates that the overall volume of the maize rhizosphere is greater than what has been supposed so far.     

Interactions between tree seedling roots and humus forms in the control of soil C and N cycling

The hypothesis that roots enhance soil-N turnover in humified soil organic matter (SOM) (mull) but not in lignified SOM (mor) was tested in a study involving the growth of eight species of tree seedlings on the two contrasting humus forms. After 12 and 22 weeks of seedling growth, soil-CO₂ efflux was measured with (1) growing seedlings, and after 22 weeks, with (2) roots only, shoots excised, and (3) with roots removed and soils amended with different rates of glucose. Indices of C-flux and of soil available-C were derived and compared to plant-N uptake, extractable soil mineral-N, anaerobically mineralized soil-N, N bioavailability to Agrostis grass following harvest of seedlings, and to seedling fine root C-chemistry. Significant soil x species interactions were found for total soil-CO₂ efflux, root-dependent CO₂, soil available-C and microbial biomass. In all cases, roots were important contributors to C-cycling in the mull soil but not in the mor soil. C was more limiting in the mor than in the mull microbial community. Plant-N uptake and the mineral-N pool was greater in the mor soil, reflecting that soil's higher specific N-supplying capacity (N-mineralized:CO₂). Seedlings decreased the mineral-N pool in both soils, but the presence of roots increased N-mineralization in the mull soil and decreased N-mineralization in the mor soil. Significant positive relationships were observed in the mull soil only between soil respiration and plant N uptake at mid-season, and between soil respiration and N-mineralization at late-season. Birch root activity in the mull soil was greater than that of all other seedlings and this observation is discussed with respect to the autecology of birch. Soil respiration correlated with the non-polar extract content but not the lignin:N ratio of fine roots. Results suggest that root-released C in mull SOM is sufficient to relieve energy limitation to soil microbes and allow them to access appreciable amounts of soil-N, whereas ligninolytic activity, which may ultimately control soil-N turnover in mor SOM, is not increased by rhizodeposition.     

Climate change goes underground: effects of elevated atmospheric CO<Subscript>2</Subscript> on microbial community structure and activities in the rhizosphere

General concern about climate change has led to growing interest in the responses of terrestrial ecosystems to elevated concentrations of CO₂ in the atmosphere. Experimentation during the last two to three decades using a large variety of approaches has provided sufficient information to conclude that enrichment of atmospheric CO₂ may have severe impact on terrestrial ecosystems. This impact is mainly due to the changes in the organic C dynamics as a result of the effects of elevated CO₂ on the primary source of organic C in soil, i.e., plant photosynthesis. As the majority of life in soil is heterotrophic and dependent on the input of plant-derived organic C, the activity and functioning of soil organisms will greatly be influenced by changes in the atmospheric CO₂ concentration. In this review, we examine the current state of the art with respect to effects of elevated atmospheric CO₂ on soil microbial communities, with a focus on microbial community structure. On the basis of the existing information, we conclude that the main effects of elevated atmospheric CO₂ on soil microbiota occur via plant metabolism and root secretion, especially in C3 plants, thereby directly affecting the mycorrhizal, bacterial, and fungal communities in the close vicinity of the root. There is little or no direct effect on the microbial community of the bulk soil. In particular, we have explored the impact of these changes on rhizosphere interactions and ecosystem processes, including food web interactions.     

Contribution of rhizomicrobial and root respiration to the CO2 emission from soil (A review)

Separate determination of root respiration and rhizomicrobial respiration is one of the most interesting, important, and methodologically complicated problems in the study of the carbon budget in soils and the subdivision of the CO₂ emission from soils into separate fluxes. In this review, we compare the main principles, the advantages and disadvantages, and the results obtained by the methods of component integration, substrate-induced respiration, respiratory capacity, girdling, isotope dilution, model rhizodeposition, modeling of the ¹⁴CO₂ efflux dynamics, exudates elution, and the δ¹³C measurements of the microbial biomass and CO₂. Summarizing the results of the determinations performed by these methods, we argue that about 40% of the rhizosphere CO₂ efflux is due to root respiration and about 60% of this efflux is due to the respiration of microorganisms decomposing root exudates.     

Three‐source partitioning of CO2 efflux from maize field soil by 13C natural abundance

A natural‐¹³C‐labeling approach—formerly observed under controlled conditions—was tested in the field to partition total soil CO₂ efflux into root respiration, rhizomicrobial respiration, and soil organic matter (SOM) decomposition. Different results were expected in the field due to different climate, site, and microbial properties in contrast to the laboratory. Within this isotopic method, maize was planted on soil with C₃‐vegetation history and the total CO₂ efflux from soil was subdivided by isotopic mass balance. The C₄‐derived C in soil microbial biomass was also determined. Additionally, in a root‐exclusion approach, root‐ and SOM‐derived CO₂ were determined by the total CO₂ effluxes from maize (Zea mays L.) and bare‐fallow plots. In both approaches, maize‐derived CO₂ contributed 22% to 35% to the total CO₂ efflux during the growth period, which was comparable to other field studies. In our laboratory study, this CO₂ fraction was tripled due to different climate, soil, and sampling conditions. In the natural‐¹³C‐labeling approach, rhizomicrobial respiration was low compared to other studies, which was related to a low amount of C₄‐derived microbial biomass. At the end of the growth period, however, 64% root respiration and 36% rhizomicrobial respiration in relation to total root‐derived CO₂ were calculated when considering high isotopic fractionations between SOM, microbial biomass, and CO₂. This relationship was closer to the 50% : 50% partitioning described in the literature than without fractionation (23% root respiration, 77% rhizomicrobial respiration). Fractionation processes of ¹³C must be taken into account when calculating CO₂ partitioning in soil. Both methods—natural ¹³C labeling and root exclusion—showed the same partitioning results when ¹³C isotopic fractionation during microbial respiration was considered and may therefore be used to separate plant‐ and SOM‐derived CO₂ sources.     

Soil nitrogen availability alters rhizodeposition carbon flux into the soil microbial community

Purpose

Soil microorganisms are important in the cycling of plant nutrients. Soil microbial biomass, community structure, and activity are mainly affected by carbon substrate and nutrient availability. The objective was to test if both the overall soil microbial community structure and the community-utilizing plant-derived carbon entering the soil as rhizodeposition were affected by soil carbon (C) and nitrogen (N) availability.

Materials and methods

A ¹³C-CO₂ steady-state labeling experiment was conducted in a ryegrass system. Four soil treatments were established: control, amendment with carboxymethyl cellulose (CMC), amendment with ammonium nitrate (NF), combined CMC and NF. Soil phospholipid fatty acid (PLFA) and ¹³C labeling PLFA were extracted and detected by isotope ratio mass spectrometer.

Results and discussion

The combined CMC and NF treatment with appropriate C/N ratio (20) significantly enhanced soil microbial biomass C and N, but resulted in lower soil inorganic N concentrations. There was no significant difference in soil PLFA profile pattern between different treatments. In contrast, most of the ¹³C was distributed into PLFAs 18:2ω6,9c, 18:1ω7c, and 18:1ω9c, indicative of fungi and gram-negative bacteria. The inorganic-only treatment was distinct in ¹³C PLFA pattern from the other treatments in the first period of labeling. Factor loadings of individual PLFAs confirmed that gram-positive bacteria had relatively greater plant-derived C contents in the inorganic-only treatment, but fungi were more enriched in the other treatments.

Conclusions

Amendments with CMC can improve N transformation processes, and the ryegrass rhizodeposition carbon flux into the soil microbial community is strongly modified by soil N availability.

     

植物根际沉积与土壤微生物关系研究进展

【目的】活跃的根际微生物被喻为植物的第二套基因组,在植物的生长发育过程中发挥着关键作用。植物通过根际碳沉积影响根际土壤微生物群落的结构和功能;作为根际微生态系统中的物质流、能量流和信息流,根际碳沉积是连接大气、植物和土壤系统物质循环的重要纽带;因此,理解根际碳沉积在根际微生态中的作用对于提高植物抗逆性,增加作物产量,调控根际养分循环等方面具有重大的理论意义。【主要进展】本文就近年来关于根际微生物领域的研究成果,重点综述了根际微生物多样性和组学研究;根际碳沉积的组成和产生机理;根际微生物群落结构的形成机制;根际微生物在促进作物养分吸收、提高作物抗逆性等方面的生态功能;以及气候变化和长期施肥对植物-微生物互作关系的影响。在此基础上我们提出了未来可能的研究重点和发展方向:1)植物根际沉积物原位收集方法和检测技术的改进和发展;2)稳定同位素探针与分子生态学技术的结合,将植物、土壤和微生物三者有机地联系起来,综合分析根际界面中微生物的活性与功能;3)高通量测序、组学技术和生物信息学等新技术的引入势必使根际微生物学研究发生革命性的变化;4)随着全球气候变化和土壤肥力改变,例如全球变暖、CO2浓度升高和长期施用化肥,根际沉积物在植物-土壤-微生物中的分配与调节机制,以及这种环境选择压力下植物如何诱导根际促生菌发挥更大作用。希望通过平衡作物与微生物之间的相互关系来实现作物的高产高效,促进农田的可持续利用。     

Responses of soil organic matter and microorganisms to freeze-thaw cycles

Soil organic matter (SOM) biomarker methods were utilized in this study to investigate the responses of fungi and bacteria to freeze-thaw cycles (FTCs) and to examine freeze-thaw-induced changes in SOM composition and substrate availability. Unamended, grass-amended, and lignin-amended soil samples were subject to 10 laboratory FTCs. Three SOM fractions (free lipids, bound lipids, and lignin-derived phenols) with distinct composition, stability and source were examined with chemolysis and biomarker Gas Chromatography/Mass Spectrometry methods and the soil microbial community composition was monitored by phospholipid fatty acid (PLFA) analysis. Soil microbial respiration was also measured before and during freezing and thawing, which was not closely related to microbial biomass in the soil but more strongly controlled by substrate availability and quality. Enhanced microbial mineralization (CO₂ flush), considered to be derived from the freeze-thaw-induced release of easily decomposable organic matter from microbial cell lyses, was detected but quickly diminished with successive FTCs. The biomarker distribution demonstrated that free lipids underwent a considerable size of decrease after repeated FTCs, while bound lipids and lignin compounds remained stable. This observation indicates that labile SOM may be most influenced by increased FTCs and that free lipids may contribute indirectly to the freeze-thaw-induced CO₂ flush from the soil. PLFA analysis revealed that fungal biomass was greatly reduced while bacteria were unaffected through the lab-simulated FTCs. Microbial community shifts may be caused by freezing stress and competition for freeze-thaw-induced substrate release. This novel finding may have an impact on carbon and nutrient turnover with predicted increases in FTCs in certain areas, because fungi and bacteria have different degradation patterns of SOM and the fungi-dominated soil community is considered to have a higher carbon storage capacity than a bacteria-dominated community.     

Assessment of 10 years of CO2 fumigation on soil microbial communities and function in a sweetgum plantation

Increased vegetative growth and soil carbon (C) storage under elevated carbon dioxide concentration ([CO₂]) has been demonstrated in a number of experiments. However, the ability of ecosystems, either above- or belowground, to maintain increased C storage relies on the response of soil processes, such as those that control nitrogen (N) mineralization, to climatic change. These soil processes are mediated by microbial communities whose activity and structure may also respond to increasing atmospheric [CO₂]. We took advantage of a long-term (ca 10 y) CO₂ enrichment experiment in a sweetgum plantation located in the southeastern United States to test the hypothesis that observed increases in root production in elevated relative to ambient CO₂ plots would alter microbial community structure, increase microbial activity, and increase soil nutrient cycling. We found that elevated [CO₂] had no detectable effect on microbial community structure using 16S rRNA gene clone libraries, on microbial activity measured with extracellular enzyme activity, or on potential soil N mineralization and nitrification rates. These results support findings at other forested Free Air [CO₂] Enrichment (FACE) sites.     

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.     

Relating the biological stability of soil organic matter to energy availability in deep tropical soil profiles

Tropical subsoils contain large reservoirs of carbon (C), most of which is stored in soil organic matter (SOM). Subsoil OM is thought to be particularly stable against microbial decomposition due to various mechanisms and its position in the soil profile, potentially representing a long-term C sink. However, few experiments have explicitly investigated SOM stability and microbial activity across several orders of magnitude of soil C concentrations as a function of soil depth. The objective of this study was to evaluate the biological stability of SOM in the upper 1.4 m of tropical forest soil profiles. We did so by measuring CO₂ evolution during a 90-day laboratory incubation experiment on a sample set that was previously characterized for C and nutrient concentrations and microbial biomass. We concurrently measured the energy content of SOM using differential scanning calorimetry (DSC) as an index of the energy available for microbial metabolism, with the hypothesis that the biological stability of SOM would be inversely related to the energy contained within it. Cumulative CO₂ evolution, mean respiration rates, and the energy density of SOM (energy released during combustion normalized to soil C) all declined with soil depth (P < 0.01). Biological indices of C stability were well correlated with measures of SOM energy. There was no change in the mean respiration rate as a function of depth when normalized to soil C, and a trend toward increased respiration per-unit microbial biomass (P = 0.07). While reduced microbial respiration in subsoils suggests an increase in the biological stability of SOM, we suggest this is driven principally by concurrent declines in energy availability as measured by DSC and the size of the microbial biomass pool. On a per-unit biomass basis, subsoil OM may be as prone to decomposition and destabilization as surface SOM.