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.     

The fate of soyabean photosynthetic carbon varies in Mollisols differing in organic carbon

The fate of photosynthetically‐fixed carbon (C) in the plant–soil–microbe continuum has received much interest because of its relevance to soil C and the global C cycle. However, information on the flow of this plant C below ground and its contribution to soil C sequestration in soils with contrasting organic C (C_org) is limited. In this study, soyabean (Glycine max L. Merr.) was grown in three Mollisols with low (1.04%), medium (2.90%) and high (5.05%) C_org, respectively. Plants were labelled with ¹³CO₂ to trace the photosynthetic C dynamics in the plant–soil system for up to 288 hours. The total amount of net fixed ¹³C by plants ranged from 66 to 78 mg pot^?1, and there was no difference between soils. The amount of ¹³C in soil organic matter (SOM) increased from 1.9 to 6.1 mg pot^?1 over time in the high‐C_org soil, while it showed a non‐significant change with 2.2 mg pot^?1 (on average) in the medium‐C_org soil, and decreased from 2.9 to 0.1 mg pot^?1 in the low‐C_org soil. In the low‐C_org soil, the amount of ¹³C in soil microbes decreased markedly over time, showing a fast turnover, and had a significant correlation (P ≤ 0.01) with ¹³C in the SOM pool. However, such a relationship was not significant in the soil with high or medium C_org. These results indicate that most of the root‐derived C in the low‐C_org soil is degraded quickly by microbial activity, while the greater input of the photosynthetic C to SOM in the high‐ and/or medium‐C_org soil can probably be attributed to physical sorption of root‐derived C by SOM and minerals, thus protecting it against microbial decomposition.     

Carbon dynamics determined by natural C abundance in microcosm experiments with soils from long-term maize and rye monocultures

Understanding carbon dynamics in soil is the key to managing soil organic matter. Our objective was to quantify the carbon dynamics in microcosm experiments with soils from long-term rye and maize monocultures using natural ¹³C abundance. Microcosms with undisturbed soil columns from the surface soil (0-25 cm) and subsoil (25-50 cm) of plots cultivated with rye (C₃-plant) since 1878 and maize (C₄-plant) since 1961 with and without NPK fertilization from the long-term experiment ‘Ewiger Roggen’ in Halle, Germany, were incubated for 230 days at 8 °C and irrigated with 2 mm 10⁻² M CaCl₂ per day. Younger, C₄-derived and older, C₃-derived percentages of soil organic carbon (SOC), dissolved organic carbon (DOC), microbial biomass (C_mic) and CO₂ from heterothropic respiration were determined by natural ¹³C abundance. The percentage of maize-derived carbon was highest in CO₂ (42-79%), followed by C_mic (23-46%), DOC (5-30%) and SOC (5-14%) in the surface soils and subsoils of the maize plots. The percentage of maize-derived C was higher for the NPK plot than for the unfertilized plot and higher for the surface soils than for the subsoils. Specific production rates of DOC, CO₂-C and C_mic from the maize-derived SOC were 0.06-0.08% for DOC, 1.6-2.6% for CO₂-C and 1.9-2.7% for C_mic, respectively, and specific production rates from rye-derived SOC of the continuous maize plot were 0.03-0.05% for DOC, 0.1-0.2% for CO₂-C and 0.3-0.5% for C_mic. NPK fertilization did not affect the specific production rates. Strong correlations were found between C₄-derived C_mic and C₄-derived SOC, DOC and CO₂-C (r≥0.90), whereas the relationship between C₃-derived C_mic and C₃-derived SOC, DOC and CO₂-C was not as pronounced (r≤0.67). The results stress the different importance of former (older than 40 years) and recent (younger than 40 years) litter C inputs for the formation of different C pools in the soil.     

Turnover of soil organic matter and of microbial biomass under C3-C4 vegetation change: Consideration of C fractionation and preferential substrate utilization

Two processes contribute to changes of the δ¹³C signature in soil pools: ¹³C fractionation per se and preferential microbial utilization of various substrates with different δ¹³C signature. These two processes were disentangled by simultaneously tracking δ¹³C in three pools - soil organic matter (SOM), microbial biomass, dissolved organic carbon (DOC) - and in CO₂ efflux during incubation of 1) soil after C₃-C₄ vegetation change, and 2) the reference C₃ soil.The study was done on the Ap horizon of a loamy Gleyic Cambisol developed under C₃ vegetation. Miscanthus giganteus - a perennial C₄ plant - was grown for 12 years, and the δ¹³C signature was used to distinguish between ‘old’ SOM (>12 years) and ‘recent’ Miscanthus-derived C (<12 years). The differences in δ¹³C signature of the three C pools and of CO₂ in the reference C₃ soil were less than 1‰, and only δ¹³C of microbial biomass was significantly different compared to other pools. Nontheless, the neglecting of isotopic fractionation can cause up to 10% of errors in calculations. In contrast to the reference soil, the δ¹³C of all pools in the soil after C₃-C₄ vegetation change was significantly different. Old C contributed only 20% to the microbial biomass but 60% to CO₂. This indicates that most of the old C was decomposed by microorganisms catabolically, without being utilized for growth. Based on δ¹³C changes in DOC, CO₂ and microbial biomass during 54 days of incubation in Miscanthus and reference soils, we concluded that the main process contributing to changes of the δ¹³C signature in soil pools was preferential utilization of recent versus old C (causing an up to 9.1‰ shift in δ¹³C values) and not ¹³C fractionation per se.Based on the δ¹³C changes in SOM, we showed that the estimated turnover time of old SOM increased by two years per year in 9 years after the vegetation change. The relative increase in the turnover rate of recent microbial C was 3 times faster than that of old C indicating preferential utilization of available recent C versus the old C.Combining long-term field observations with soil incubation reveals that the turnover time of C in microbial biomass was 200 times faster than in total SOM. Our study clearly showed that estimating the residence time of easily degradable microbial compounds and biomarkers should be done at time scales reflecting microbial turnover times (days) and not those of bulk SOM turnover (years and decades). This is necessary because the absence of C reutilization is a prerequisite for correct estimation of SOM turnover. We conclude that comparing the δ¹³C signature of linked pools helps calculate the relative turnover of old and recent pools.     

Sources and mechanisms of priming effect induced in two grassland soils amended with slurry and sugar

The mechanisms and specific sources of priming effects, i.e. short term changes of soil organic matter (SOM) decomposition after substance addition, are still not fully understood. These uncertainties are partly method related, i.e. until now only two C sources in released CO₂ could be identified. We used a novel approach separating three carbon (C) sources in CO₂ efflux from soil. The approach is based on combination of different substances originated from C₃ or C₄ plants in different treatments and identical transformation of substances like C₃ sugar (from sugar beet) and C₄ sugar (from sugar cane). We investigated the influence of the addition of two substances having different microbial utilizability, i.e. slurry and sugar on the SOM or/and slurry decomposition in two grassland soils with different levels of C_org (2.3 vs. 5.1% C). Application of slurry to the soil slightly accelerated the SOM decomposition. Addition of sugar lead to changes of SOM and slurry decomposition clearly characterized by two phases: immediately after sugar addition, the microorganisms switched from the decomposition of hardly utilizable SOM to the decomposition of easily utilizable sugar. This first phase was very short (2-3 days), hence was frequently missed in other experiments. The second phase showed a slightly increased slurry and SOM decomposition (compared to the soil without sugar). The separation of three sources in CO₂ efflux from grassland soils allowed us to conclude that the C will be utilized according to its utilizability: sugar>slurry>SOM. Additionally, decomposition of more inert C (here SOM) during the period of intensive sugar decomposition was strongly reduced (negative priming effect). We conclude that, priming effects involve a chain of mechanisms: (i) preferential substrate utilization, (ii) activation of microbial biomass by easily utilizable substrate (iii) subsequent increased utilization of following substrates according to their utilizability, and (iv) decline to initial state.     

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.     

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.     

Indications for soil organic matter quality in soils under different management

Changes in management practice are reflected by soil carbon and nitrogen status, in particular by the proportion of soil organic matter (SOM) being easily transformed (active SOM). We describe SOM quality for three management practices, Organic Farming system (OF), Integrated Crop Production (ICP) and pasture sites (G), which intend to achieve sustainable management practice. The experimental sites were conventionally farmed until 1992. SOM quality was examined by describing active SOM pools, such as the decomposed ‘young soil organic matter’ (YSOM), ratio of microbial biomass carbon (C_mic) to organic carbon (C_org), ecophysiological status of the microbial biomass (qCO₂), and the ratio of light particulate organic matter (POM-LF) to C_org. Ratios of soil microbial biomass (C_mic/C_org) and POM-LF (POM-LF/C_org) and the amount of decomposed YSOM were relatively similar to each other, despite differences in management practice and soil texture. Soil microbial parameters (C_mic, C_mic/C_org and qCO₂) were significantly (p<0.05) affected by the amount of decomposed YSOM and the silt content in the OF. In the ICP, soil microbial parameters depended only on the amount of decomposed YSOM, which was considered to be a consequence of the more heterogeneous texture at the OF-sites. Management effects were detectable for no-tillage in the ICP leading to an accumulation of active SOM in the surface soil (0–10 cm). The ratio POM-LF/C_org showed no difference between G and OF despite markedly higher C_org-contents at the G-sites. Conclusively, all methods used indicate comparable SOM qualities for the three management systems, despite differences in soil texture and soil management during 7 years. Management practices seem to be well adapted to the site conditions.     

Thermal stability of soil organic matter pools and their δC values after C3-C4 vegetation change

Carbon isotopic composition of soils subjected to C₃-C₄ vegetation change is a suitable tool for the estimation of C turnover in soil organic matter (SOM) pools. We hypothesized that the biological availability of SOM pools is inversely proportional to their thermal stability. Soil samples from a field plot with 10.5 years of cultivation of the C₄ plant Miscanthus×gigantheus and from a reference plot under C₃ grassland vegetation were analysed by thermogravimetry coupled with differential scanning calorimetry (TG-DSC). According to differential weight losses (dTG) and energy release or consumption (DSC), five SOM pools with increasing thermal stability were distinguished: (I) 20-190 °C, (II) 190-310 °C, (III) 310-390 °C, (IV) 390-480 °C, and (V) 480-1000 °C. Their δ¹³C values were analysed by EA-IRMS. The weight losses in pool I were connected with water evaporation, since no significant C losses were measured and δ¹³C values remained unchanged. The δ¹³C of pools II and III in soil samples under Miscanthus were closer to the δ¹³C of the Miscanthus plant tissues (−11.8‰) compared to the thermally stable SOM pool V (−19.5‰). The portion of the Miscanthus-derived C₄-C in total SOM in 0-5 cm reached 55.4% in the 10.5 years. The C₄-C contribution in pool II was 60% and decreased down to 6% in pool V. The mean residence times (MRT) of SOM pools II, III, and IV were similar (11.6, 12.2, and 15.4 years, respectively), while pool V had a MRT of 163 years. Therefore, we concluded that the biological availability of thermal labile SOM pools (<480 °C) was higher, than that of the thermal stable pool decomposed above 480 °C. However, the increase of SOM stability with rising temperature was not gradual. Therefore, the applicability of the TG-DSC for the separation of SOM pools with different biological availability is limited.     

Effect of lithological substrate on microbial biomass and enzyme activity in brown soil profiles in the northern Apennines (Italy)

The aim of the present study was to investigate the microbial activity along forest brown soil profiles sequence developed on different lithological substrates (carbonate or non-carbonated cement in sandstone formations) at different altitudes. The main question posed was: does carbonate affect the biochemical activity of brown soil profiles at different altitudes? For the purpose of this study, four soil profiles with different amounts and compositions of SOM developed on different lithological substrates were selected: two with carbonate (MB and MZ) and the other two with non-carbonated cement in the sandstone formations (MF1 and MF2). Chemical and biochemical properties of soil were analysed along soil profiles in order to assess the SOM quantity and quality, namely total organic C (C_org), water extractable organic C (WEOC) and humification indices (HI, DH, HR). Microbial biomass (C_mic and N_mic) content, as well as the specific activities of acid phosphatase, β-glucosidase and chitinase enzymes were chosen as indicators of biochemical activity. The soil biochemical properties provided evidence of better conditions for microorganisms in MB than in MF1, MF2 and MZ soil profiles, since patterns of microbial biomass content and activity might be expected in response to the amount and quality of organic substances. The different lithological substrates did not show any clear effect on soil microbial biomass content, since similar values were obtained in MF1, MF2 (with non-carbonated cement) and MZ (with carbonate). However, the specific activities of acid phosphatase (per unit of C_org and per unit of C_mic) were higher in soils with no carbonate (MF1 and MF2) than in soils with carbonate (MB and MZ). In conclusion, the biochemical activity along brown soil profiles was mainly regulated by different soil organic matter content and quality, while the two different lithological substrates (with carbonate or non-carbonated cement in the sandstone formations) did not show any direct effect on microbial biomass and its activity. However, the activity of acid phosphatase per unit of C was particularly enhanced in soil with non-carbonate cement in the sandstone formations.     

Changes in forest soil organic matter pools after a decade of elevated CO2 and O3

The impact of rising atmospheric carbon dioxide (CO₂) may be mitigated, in part, by enhanced rates of net primary production and greater C storage in plant biomass and soil organic matter (SOM). However, C sequestration in forest soils may be offset by other environmental changes such as increasing tropospheric ozone (O₃) or vary based on species-specific growth responses to elevated CO₂. To understand how projected increases in atmospheric CO₂ and O₃ alter SOM formation, we used physical fractionation to characterize soil C and N at the Rhinelander Free Air CO₂-O₃ Enrichment (FACE) experiment. Tracer amounts of ¹⁵NH₄⁺ were applied to the forest floor of Populus tremuloides, P. tremuloides-Betula papyrifera and P. tremuloides-Acer saccharum communities exposed to factorial CO₂ and O₃ treatments. The ¹⁵N tracer and strongly depleted ¹³C-CO₂ were traced into SOM fractions over four years. Over time, C and N increased in coarse particulate organic matter (cPOM) and decreased in mineral-associated organic matter (MAOM) under elevated CO₂ relative to ambient CO₂. As main effects, neither CO₂ nor O₃ significantly altered ¹⁵N recovery in SOM. Elevated CO₂ significantly increased new C in all SOM fractions, and significantly decreased old C in fine POM (fPOM) and MAOM over the duration of our study. Overall, our observations indicate that elevated CO₂ has altered SOM cycling at this site to favor C and N accumulation in less stable pools, with more rapid turnover. Elevated O₃ had the opposite effect, significantly reducing cPOM N by 15% and significantly increasing the C:N ratio by 7%. Our results demonstrate that CO₂ can enhance SOM turnover, potentially limiting long-term C sequestration in terrestrial ecosystems; plant community composition is an important determinant of the magnitude of this response.     

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.     

Spatial variability in the carbon of microbial biomass and microbial respiration in soils of the south of Moscow oblast

Soil samples from the upper 10-cm-thick layer of the humus horizon (without forest litter) were taken in Podol’sk and Serpukhov districts (1130 and 1080 km², respectively) of Moscow oblast. At each sampling site, ecosystem (forest, plowland, or fallow), soil (soddy-podzolic, soddy-gley, bog-podzolic, meadow alluvial, gray forest, and anthropogenically transformed soils of lawns and industrial zones), predominant vegetation, and topography (floodplain and low, medium, and upper parts of watersheds) were determined. The carbon content of the microbial biomass (C_mic) was determined by the method of substrate-induced respiration; we also determined the rate of basal (microbial) respiration (BR) and the organic carbon content, pH, and particle-size distribution. Overall, 237 samples from Serpukhov district and 45 samples from Podol’sk district were analyzed. The BR/C_mic ratios (respiration quotient qCO₂) and C_mic/C_org ratios were calculated. The C_mic content in the soils ranged from 43 to 1394 μg C/kg; the BR varied from 0.06 to 25 μg CO₂-C/g per h, qCO₂, from 0.34 to 6.52 μg CO₂-C/mg C_mic per h; and the C_mic/C_org ratio, from 0.19 to 10.65%. It was found that the most significant factors affecting the variability of the C_mic and BR are the parameters of ecosystem (50% and 80%, respectively) and soil (30% and 9%, respectively). The most significant variability of these indices was found in forest soils; it was mainly controlled by the soil texture (33 and 23%) and the C_org content (19 and 24%). The C_mic parameter made it possible to differentiate the soils of the territory for the purposes of their evaluation, monitoring, and biological assessment more clearly than the BR value and the soil chemical characteristics.     

The fate of catechol in soil as affected by earthworms and clay

The effect of endogeic earthworms (Octolasion tyrtaeum) and the availability of clay (Montmorillonite) on the mobilization and stabilization of uniformly ¹⁴C-labelled catechol mixed into arable and forest soil was investigated in a short- and a long-term microcosm experiment. By using arable and forest soil the effect of earthworms and clay in soils differing in the saturation of the mineral matrix with organic matter was investigated. In the short-term experiment microcosms were destructively sampled when the soil had been transformed into casts. In the long-term experiment earthworm casts produced during 7 days and non-processed soil were incubated for three further months. Production of CO₂ and ¹⁴CO₂ were measured at regular intervals. Accumulation of ¹⁴C in humic fractions (DOM, fulvic acids, humic acids and humin) of the casts and the non-processed soil and incorporation of ¹⁴C into earthworm tissue were determined.Incorporation of ¹⁴C into earthworm tissue was low, with 0.1 and 0.44% recovered in the short- and long-term experiment, respectively, suggesting that endogeic earthworms preferentially assimilate non-phenolic soil carbon. Cumulative production of CO₂-C was significantly increased in casts produced from the arable soil, but lower in casts produced from the forest soil; generally, the production of CO₂-C was higher in forest than in arable soil. Both soils differed in the pattern of ¹⁴CO₂-C production; initially it was higher in the forest soil than in the arable soil, whereas later the opposite was true. Octolasion tyrtaeum did not affect ¹⁴CO₂-C production in the forest soil, but increased it in the arable soil early in the experiment; clay counteracted this effect. Clay and O. tyrtaeum did not affect integration of ¹⁴C into humic fractions of the forest soil. In contrast, in the arable soil O. tyrtaeum increased the amount of ¹⁴C in the labile fractions, whereas clay increased it in the humin fraction.The results indicate that endogeic earthworms increase microbial activity and thus mineralization of phenolic compounds, whereas clay decreases it presumably by binding phenolic compounds to clay particles when passing through the earthworm gut. Endogeic earthworms and clay are only of minor importance for the fate of catechol in soils with high organic matter, clay and microbial biomass concentrations, but in contrast affect the fate of phenolic compounds in low clay soils.     

Determination of the fate of regularly applied 13C‐labeled‐artificial‐exudates C in two agricultural soils

Exudates are part of the total rhizodeposition released by plant roots to soil and are considered as a substantial input of soil organic matter. Exact quantitative data concerning the contribution of exudates to soil C pools are still missing. This study was conducted to reveal effects of ¹³C‐labeled exudate (artificial mixture) which was regularly applied to upper soil material from two agricultural soils. The contribution of exudate C to water‐extractable organic C (WEOC), microbial biomass C (MBC), and CO₂‐C evolution was investigated during a 74 d incubation. The WEOC, MBC, and CO₂‐C concentrations and the respective δ¹³C values were determined regularly. In both soils, significant incorporation of artificial‐exudate‐derived C was observed in the WEOC and MBC pool and in CO₂‐C. Up to approx. 50% of the exudate‐C amounts added were recovered in the order WEOC << MBC < CO₂‐C in both soils at the end of the incubation. Newly built microbial biomass consisted mainly of exudates, which substituted soil‐derived C. Correspondingly, the CO₂‐C evolved from exudate‐treated soils relative to the controls was dominated by exudate C, showing a preferential mineralization of this substrate. Our results suggest that the remaining 50% of the exudate C added became stabilized in non‐water‐extractable organic fractions. This assumption was supported by the determination of the total organic C in the soils on the second‐last sampling towards the end of the incubation. In the exudate‐treated soils, significantly more soil‐derived C compared to the controls was found in the WEOC on almost all samplings and in the MBC on the first sampling. This material might have derived from exchange processes between the added exudate and the soil matrix. This study showed that easily available substrates can be stabilized in soil at least in the short term.     

Three-source partitioning of CO2 efflux from soil planted with maize by C natural abundance fails due to inactive microbial biomass

A theoretical approach to the partitioning of carbon dioxide (CO₂) efflux from soil with a C₃ vegetation history planted with maize (Zea mays), a C₄ plant, into three sources, root respiration (RR), rhizomicrobial respiration (RMR), and microbial soil organic matter (SOM) decomposition (SOMD), was examined. The δ¹³C values of SOM, roots, microbial biomass, and total CO₂ efflux were measured during a 40-day growing period. A three-source isotopic mass balance based on the measured δ¹³C values and on assumptions made in other studies showed that RR, RMR, and SOMD amounted to 91%, 4%, and 5%, respectively. Two assumptions were thoroughly examined in a sensitivity analysis: the absence of ¹³C fractionation and the conformity of δ¹³C of microbial CO₂ and that of microbial biomass. This approach strongly overestimated RR and underestimated RMR and microbial SOMD. CO₂ efflux from unplanted soil was enriched in ¹³C by 2.0‰ compared to microbial biomass. The consideration of this ¹³C fractionation in the mass balance equation changed the proportions of RR and RMR by only 4% and did not affect SOMD. A calculated δ¹³C value of microbial CO₂ by a mass balance equation including active and inactive parts of microbial biomass was used to adjust a hypothetical below-ground CO₂ partitioning to the measured and literature data. The active microbial biomass in the rhizosphere amounted to 37% to achieve an appropriate ratio between RR and RMR compared to measured data. Therefore, the three-source partitioning approach failed due to a low active portion of microbial biomass, which is the main microbial CO₂ source controlling the δ¹³C value of total microbial biomass. Since fumigation-extraction reflects total microbial biomass, its δ¹³C value was unsuitable to predict δ¹³C of released microbial CO₂ after a C₃-C₄ vegetation change. The second adjustment to the CO₂ partitioning results in the literature showed that at least 71% of the active microbial biomass utilizing maize rhizodeposits would be necessary to achieve that proportion between RR and RMR observed by other approaches based on ¹⁴C labelling. The method for partitioning total below-ground CO₂ efflux into three sources using a natural ¹³C labelling technique failed due to the small proportion of active microbial biomass in the rhizosphere. This small active fraction led to a discrepancy between δ¹³C values of microbial biomass and of microbially respired CO₂.     

C and N turnover in structurally intact soils of different texture

The turnover of native and applied C and N in undisturbed soil samples of different texture but similar mineralogical composition, origin and cropping history was evaluated at −10 kPa water potential. Cores of structurally intact soil with 108, 224 and 337 g clay kg⁻¹ were horizontially sliced and ¹⁵N-labelled sheep faeces was placed between the two halves of the intact core. The cores together with unamended treatments were incubated in the dark at 20 °C and the evolution of CO₂-C determined continuously for 177 d. Inorganic and microbial biomass N and ¹⁵N were determined periodically. Net nitrification was less in soil amended with faeces compared with unamended soil. When adjusted for the NO₃-N present in soil before faeces was applied, net nitrification became negative indicating that NO₃-N had been immobilized or denitrified. The soil most rich in clay nitrified least N and ¹⁵N. The amounts of N retained in the microbial biomass in unamended soils increased with clay content. A maximum of 13% of the faeces ¹⁵N was recovered in the microbial biomass in the amended soils. CO₂-C evolution increased with clay content in amended and unamended soils. CO₂-C evolution from the most sandy soil was reduced due to a low content of potentially mineralizable native soil C whereas the rate constant of C mineralization rate peaked in this soil. When the pool of potentially mineralizable native soil C was assumed proportional to volumetric water content, the three soils contained similar proportions of potentially mineralizable native soil C but the rate constant of C mineralization remained highest in the soil with least clay. Thus although a similar availability of water in the three soils was ensured by their identical matric potential, the actual volume of water seemed to determine the proportion of total C that was potentially mineralizable. The proportion of mineralizable C in the faeces was similar in the three soils (70% of total C), again with a higher rate constant of C mineralization in the soil with least clay. It is hypothesized that the pool of potentially mineralizable C and C rate constants fluctuate with the soil water content.     

Specific features of the ecological functioning of urban soils in Moscow and Moscow oblast

Urban soils (constructozems) were studied in Moscow and several cities (Dubna, Pushchino, and Serebryanye Prudy) of Moscow oblast. The soil sampling from the upper 10-cm-thick layer was performed in the industrial, residential, and recreational functional zones of these cities. The biological (the carbon of the microbial biomass carbon, C_mic and the microbial (basal) respiration, BR) and chemical (pH_water and the contents of C_org, heavy metals, and NPK) indices were determined in the samples. The ratios of BR to C_mic (the microbial respiration quotient, qCO₂) and of C_mic to C_org were calculated. The C_mic varied from 120 to 738 μg C/g soil; the BR, from 0.39 to 1.94 μg CO₂-C/g soil per hour; the C_org, from 2.52 to 5.67%; the qCO₂, from 1.24 to 5.28 μg CO₂-C/mg C_mic/g soil per h; and the C_mic/C_org, from 0.40 to 1.55%. Reliable positive correlations were found between the C_mic and BR, the C_mic and C_mic/C_org, and the C_mic and C_org values (r = 0.75, 0.95, and 0.61, respectively), as well as between the BR and C_mic/C_org values (r = 0.68). The correlation between the C_mic/C_org and qCO₂ values was negative (r = −0.70). The values of C_mic, BR, C_org, and C_mic/C_org were found to correlate with the ammonium nitrogen content. No correlative relationships were revealed between the determined indices and the climatic characteristics. The principal component analysis described 86% of the variances for all the experimental data and clearly subdivided the locations of the studied soil objects. The ANOVA showed that the variances of C_mic, C_org, and BR are controlled by the site location factor by 66, 63, and 35%, respectively. The specificity of the functioning of the anthropogenic soils as compared with their natural analogues was clearly demonstrated. As shown in this study, measurable biological indices might be applied to characterize the ecological, environmental-regulating, and productive functions of soils, including urban soils.     

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.     

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.