Effect of barley plants on the decomposition of 14C-labelled soil organic matter

The effect of barley plants on the rate of decomposition of soil organic matter over a 6-week period was studied using soil that had been previously labelled by incubation with ¹⁴C-labelled ryegrass for 1 year. The plants reduced the loss of ¹⁴CO₂, from soil by 70 per cent over 42 days. About half of the reduction was accounted for by the uptake of labelled C by the plant roots, very little ¹⁴C label being associated with the shoot. Chemical fractionation of the root showed that the ¹⁴C was chemically incorporated into cell wall materials such as cellulose and holocellulose. The reduction in organic matter decomposition in the presence of plants has been explained by earlier workers in terms ofa reduction in microbial activity as a result of a soil moisture deficit caused by plant transpiration. This explanation does not account for all the reduction in decomposition noted in the present experiments. Control soil (without a plant, but amended with glucose or yeast extract to simulate the effect of root exudates) showed a small positive priming effect, the release of ¹⁴CO₂, being increased. Thus the mechanism by which plants conserve organic matter is complex and cannot be explained merely by analogy to an increased level of nutrients available for microbial metabolism.     

Suppression of decomposition of 14C-labelled plant roots in the presence of living roots of maize and perennial ryegrass

The rates of decomposition of barley roots labelled with ¹⁴C were investigated in soil planted with maize or perennial ryegrass and in fallow controls. Evolution of ¹⁴CO₂ was significantly less from the planted soils than from fallow controls. Roots of maize and ryegrass appeared to compete substantially with soil microbes for ¹⁴C-labelled materials. Simple competitive effects were, however, insufficient to explain all of the observed effects of root growth on soil organic matter decomposition. There was no indication that the detrimental effects of maize roots on aggregate stability could be associated with increased degradation of native soil organic materials; the broader significance of the results is also discussed.     

Contributions of incorporated residue and living roots to aggregate-associated and microbial carbon in two soils with different clay mineralogy

We conducted a study to investigate the role of aggregates in the stabilization of residue‐ and root‐derived C in an illitic Mollisol and a kaolinitic Oxisol under the following treatments: (i) incorporated residue, (ii) growing plants, and (iii) both incorporated residue and growing plants. Residue‐C dynamics were followed in soils incubated with ¹³C‐labelled wheat residue with and without unlabelled growing wheat plants. Root‐C was traced by growing wheat plants with and without unlabelled wheat residue in a ¹³CO₂‐labelling chamber. After 46 and 76 incubation days, residue‐ and root‐C were measured in four aggregate size classes and in microbial‐C. Both soils had greater residue‐derived than root‐derived total aggregate‐associated C at day 76, which we attributed to the larger residue‐C than root‐C inputs at the start of the experiment. On an aggregate basis, the ratio of residue‐derived over root‐derived C decreased in most size fractions over time, indicating a greater potential for longer‐term root‐C than residue‐C stabilization by aggregates in both soils. At both sampling days, all aggregates > 53 µm had greater residue‐C concentrations in the illitic soil than in the kaolinitic soil and this difference increased with increasing aggregate size. This suggested a greater affinity of illite clay than kaolinite clay to bind with fresh residue‐derived compounds into larger aggregates and hence a greater importance of aggregates in stabilizing residue‐C in illitic compared with kaolinitic soils. The stabilization of root‐C by aggregates was less affected by clay mineralogy and thus less dependent on the affinity of clay minerals to bind with root‐derived compounds.     

Transformation and binding of 13C and 14C-labelled atrazine in relation to straw decomposition in soil

As a source of organic matter, crop residues affect the behaviour of pesticides in agricultural soils. The fate of [U‐ring‐¹³C] and [U‐ring‐¹⁴C] atrazine (6‐chloro‐N‐ethyl‐N‐isopropyl‐1,3,5‐triazine‐2,4‐diamine) was investigated during laboratory incubation under controlled conditions in a loamy soil amended with wheat straw at two different states of decomposition: no preliminary decomposition or 6 months’ preliminary decomposition. After 3 months, non‐extractable, so‐called ‘bound’, ¹³C‐atrazine residues were recovered in three particle‐size fractions (> 200, 50–200 and < 50 μm), and investigated with solid‐state ¹³C‐NMR spectroscopy. Parallel incubations with [U‐ring‐¹⁴C] atrazine were carried out to quantify the bound residues as well as the extractable and mineralized fractions. The effect of straw residues on atrazine behaviour depended on whether they had been previously decomposed or not. When straw was decomposed for 6 months prior to incubation, atrazine mineralization was enhanced to 50% of the initial ¹⁴C in contrast to 15% of the initial ¹⁴C in soil alone and soil amended with fresh straw. In parallel, atrazine bound residues were formed in greater amount representing up to 20% of the initial ¹⁴C. CP/MAS ¹³C‐NMR on soil size fractions of soil–straw mixtures after incubation with ¹³C‐atrazine showed that bound residues contained mostly triazinic C, corresponding to atrazine or primary metabolites. Non‐humified organic materials recovered in size fractions > 200 and 50–200 μm contained significant amounts of bound residues, especially when straw was added to the soil. CP/MAS ¹³C‐NMR analysis of humic acids obtained from < 50‐μm fractions was difficult due to overlapping of the native carboxyl ¹³C signal with the ¹³C‐atrazine signal.     

Stabilization of carbon from maize in a sandy soil in a long-term experiment

The sequestration of carbon (C) in soil is not completely understood, and quantitative information about the amounts of organic carbon in the various fractions and their rates of turnover could improve understanding. We aimed (i) to quantify the amounts of C derived from maize at various depths in the soil in a long‐term field experiment with and without fertilization using ¹³C/¹²C analysis, (ii) to model changes in the organic C, and (iii) to compare measured and modelled pools of C. The organic C derived from the maize was measured in soil samples collected to a depth of 65 cm from four plots, two of which had been under continuous maize and two under continuous rye during long‐term field experiments with NPK and without fertilization. The fractionation procedures included particle‐size fractionation and extractions in water and in pyrophosphate solution. We used the Rothamsted Carbon Model to model the dynamics of the carbon from ¹³C data. The amounts of C derived from maize in the Ap horizon after 39 years of continuous maize cropping were 9.5% of the total organic C (where unfertilized) and 14.0% where NPK had been applied. Fertilization did not affect the residence time of carbon in the soil. The amounts of C derived from maize in water extracts were 21% of the total organic C (where unfertilized) and 22% where NPK had been applied. The extracts that were soluble in pyrophosphate and insoluble in acid were depleted in C from maize (the amounts were 5% and 7% of the total organic C, respectively). The results of the ¹³C natural abundance technique were used to model the dynamics of the organic C. Both the total organic C and the C derived from maize in the particle‐size fraction 0–63 μm agreed well with the total and maize‐derived sums of the model pools ‘inert organic matter’, ‘humified organic matter’ and ‘microbial biomass’. The model suggested that 64% (unfertilized) or 53% (NPK) of the organic C in the Ap horizon were inert. Only one of three published equations to determine the size of the inert pool agreed well with these model results.     

Plant rhizodeposition — an important source for carbon turnover in soils

The soil organic matter plays a key role in ecological soil functions, and has to be considered as an important CO₂ sink on a global scale. Apart from crop residues (shoots and roots), left over on the field after harvest, carbon and nitrogen compounds are also released by plant roots into the soil during vegetation, and undergo several transformation processes. Up to now the knowledge about amount, composition, and turnover of these root‐borne compounds is still very limited. So far it could be demonstrated with different plant species, that up to 20 % of photosynthetically fixed C are released into the soil during vegetation period. These C amounts are ecological relevant. Depending on assimilate sink strength during ontogenesis, the C release varies with plant age. A large percentage of these root‐borne substances were rapidly respired by microorganisms (64—86 %). About 2—5 % of net C assimilation was kept in soil. The root exudates of maize were mainly water‐soluble (79 %), and in this fraction about 64 % carbohydrates, 22 % amino acids/amides and 14 % organic acids could be identified. Plant species and in some cases also plant cultivars varied strongly in their root exudation pattern. Under non‐sterile conditions the exuded compounds were rapidly stabilized in water‐insoluble forms and bound preferably to the soil clay fraction. The binding of root exudates to soil particles also improved soil structure by increasing aggregate stability. Future research should focus on quantification and characterization of root‐borne C compounds during the whole plant ontogenesis. Apart from pot experiments with ¹⁴CO₂ labeling, it is necessary to conduct model field experiments with ¹³CO₂ labeling in order to be able to distinguish between CO₂ originating from the soil C pool and rhizosphere respiration, originating from plant assimilates. Such a separation is necessary to assess if soils are sources or sinks of CO₂. The incorporation of root‐borne C (¹⁴C, ¹³C) into soil organic matter of different stability is also of particular interest.     

The dynamics of organic matter in rock fragments in soil investigated by 14C dating and measurements of 13C

Rock fragments in soil can contain significant amounts of organic carbon. We investigated the nature and dynamics of organic matter in rock fragments in the upper horizons of a forest soil derived from sandstone and compared them with the fine earth fraction (<2 mm). The organic C content and its distribution among humic, humin and non‐humic fractions, as well as the isotopic signatures (Δ¹⁴C and δ¹³C) of organic carbon and of CO₂ produced during incubation of samples, all show that altered rock fragments contain a dynamic component of the carbon cycle. Rock fragments, especially the highly altered ones, contributed 4.5% to the total organic C content in the soil. The bulk organic matter in both fine earth and highly altered rock fragments in the A1 horizon contained significant amounts of recent C (bomb ¹⁴C), indicating that most of this C is cycled quickly in both fractions. In the A horizons, the mean residence times of humic substances from highly altered rock fragments were shorter than those of the humic substances isolated in the fine earth. Values of Δ¹⁴C of the CO₂ produced during basal respiration confirmed the heterogeneity, complexity and dynamic nature of the organic matter of these rock fragments. The weak ¹⁴C signatures of humic substances from the slightly altered rock fragments confirmed the importance of weathering in establishing and improving the interactions between rock fragments and surrounding soil. The progressive enrichment in ¹³C from components with high‐¹⁴C (more recent) to low‐¹⁴C (older) indicated that biological activity occurred in both the fine and the coarse fractions. Hence the microflora utilizes energy sources contained in all the soil compartments, and rock fragments are chemically and biologically active in soil, where they form a continuum with the fine earth.     

Effect of active roots on the decomposition of soil organic materials

Summary The effect of one form of soil organic matter, such as living roots or root exudates on another form of soil organic matter, such as dead roots or incorporated litter and litter leachates, has been studied from various perspectives over the last 25 years. The effect seems to be either positive (priming) or negative (conserving). The present review concentrates on the conserving effect, measured as a decrease in ¹⁴CO₂ released, in both field and greenhouse/growth chamber studies. The field experiments suggested that certain physical conditions in the soil, such as less available moisture or restricted aeration which led to lower microbial activity, explained the conserving effect of living roots on soil organic matter. Although more detailed greenhouse/growth chamber studies confirmed the conserving effect per se, it appears that biological rather than physical factors could better explain the reduction in the rate of decomposition of ¹⁴C-labelled plant residues in the presence of roots. However, a complex picture has emerged through a variety of postulates, all proposed in attempts to explain the conserving effect. Finally, the most recent studies have argued that the decrease in decomposition of labelled organic matter in planted soil is probably more apparent than real. A decrease in respired ¹⁴CO₂ could be explained by an incorporation of ¹⁴C derived from old roots into the rhizosphere microbial populations of the living roots. To make any further progress on the fundamental question of how soil organic matter moves along its continuum from a living to a refractory state, the microenvironment needs to be examined at periodic intervals. New developments in improved histochemical and electron-probe microanalyses look promising.LRS Contribution no. 3878970     

Determination of the fate of 13C labelled maize and wheat exudates in an agricultural soil during a short-term incubation

A broader knowledge of the contribution of carbon (C) released by plant roots (exudates) to soil is a prerequisite for optimizing the management of organic matter in arable soils. This is the first study to show the contribution of constantly applied ¹³C‐labelled maize and wheat exudates to water extractable organic carbon (WEOC), microbial biomass‐C (MB‐C), and CO₂‐C evolution during a 25‐day incubation of agricultural soil material. The CO₂‐C evolution and respective δ¹³C values were measured daily. The WEOC and MB‐C contents were determined weekly and a newly developed method for determining δ¹³C values in soil extracts was applied. Around 36% of exudate‐C of both plants was recovered after the incubation, in the order WEOC < MB‐C < CO₂‐C for maize and MB‐C < WEOC < CO₂‐C for wheat. Around 64% of added exudate‐C was not retrieved with the methods used here. Our results suggest that great amounts of exudates became stabilized in non‐water extractable organic fractions. The amounts of MB‐C stayed relatively constant over time despite a continuous exudate‐C supply, which is the prerequisite for a growing microbial population. A lack of mineral nutrients might have limited microbial growth. The CO₂‐C mineralization rate declined during the incubation and this was probably caused by a shift in the microbial community structure. Consequently, incoming WEOC was left in the soil solution leading to rising WEOC amounts over time. In the exudate‐treated soil additional amounts of soil‐derived WEOC (up to 110 μg g⁻¹) and MB‐C (up to 60 μg g⁻¹) relative to the control were determined. We suggest therefore that positive priming effects (i.e. accelerated turnover of soil organic matter due to the addition of organic substrates) can be explained by exchange processes between charged, soluble C‐components and the soil matrix. As a result of this exchange, soil‐derived WEOC becomes available for mineralization.     

Nitrogen in particle size fractions of soils incubated for five years with 15N-ammonium and 14C-hemicellulose

Four soils with a range of clay and silt contents were incubated for 5 a with ¹⁵N-labelled (NH₄)SO₄ and ¹⁴C-labelled hemicellulose and then fractionated according to particle size by ultrasonic dispersion and sedimentation. The distribution of labelled and native N between clay, silt and sand fractions was determined and elated to previous results on the C distributions. Between 29% and 48% of the added N was found in organic form. The ¹⁵N atom percentage excess decreased in the order: clay > whole soil > silt > sand. For both clay and silt, the enrichment factor for labelled and native N decreased with increasing fraction weight. Clay enrichment was higher for labelled than for native N, the converse being true for silt. The distribution of whole soil labelled organic N was: clay 77–91%, silt 4–11%, and sand <0.5%. Corresponding values for native N were 69–74%, 16–22%, and 1–2%, respectively. All soils had higher proportions of labelled than of native N in the clay, the converse was true for the silt. The C/N ratio of the native silt organic matter was higher and that of clay organic matter lower than whole soil C/N ratios. Differences between the C/N ratio distributions of native and labelled organic matter were small. The relative distribution of labelled N and C was very similar confirming that the turnover of C and N in soil organic matter is closely interrelated.     

Distribution of phosphorus in size fractions of sandy soils with different fertilization histories

A major challenge in sustainable crop management is to ensure adequate P supply for crops, while minimizing losses of P that could negatively impact water quality. The objective of the present study was to investigate the effects of long‐term applications of different levels of mineral fertilizers and farmyard manure on (1) the availability of P, (2) the relationship between soil C, N, and P, and (3) the distribution of inorganic and organic P in size fractions obtained by wet sieving. Soil samples were taken from the top 20 cm of a long‐term (29 y) fertilization trial on a sandy Cambisol near Darmstadt, SW Germany. Plant‐available P, determined with the CAL method, was little affected by fertilization treatment (p < 0.05) and was low to optimal. The concentration of inorganic and organic P extracted with a NaOH‐EDTA solution (P_NaOH‐EDTA) averaged about 350 mg (kg dry soil)^–1, with 42% being in the organic form (P_o). Manure application tended to increase soil C, N, and P_o concentrations by 8%, 9%, and 5.6%, respectively. Across all treatments, the C : N : P_o ratio was 100 : 9.5 : 2 and was not significantly affected by the fertilization treatments. Aggregate formation was weak due to the low clay and organic‐matter content of the soil, and the fractions > 53 μm consisted predominantly of sand grains. The different fertilization treatments had little effect on the distribution of size fractions and their C, N, and P contents. In the fractions > 53 μm, P_NaOH‐EDTA ranged between 200 and 300 mg kg^–1, while it reached 1260 mg kg^–1 in the fraction < 53 μm. Less than one third of P_NaOH‐EDTA was present as P_o in the fractions > 53 μm, while P_o accounted for 70% of P_NaOH‐EDTA in the smallest fraction (< 53 μm). Therefore, 16% and 28% of P_NaOH‐EDTA and P_o, respectively, were associated with the smallest fraction, even though this fraction accounted for < 5% of the soil mass. Therefore, runoff may cause higher P losses than the soil P content suggests in this sandy soil with a weak aggregate formation. Overall, the results indicate that manure and mineral fertilizer had similar effects on soil P fractions.     

A 13C tracer study to identify the origin of dissolved organic carbon in forested mineral soils

The relative contributions of sources of carbon in soils, such as throughfall, litter, roots, microbial decay products and stable organic fractions, to dissolved organic C are controversial. To identify the origin of dissolved organic C, we made use of a 4‐year experiment where spruce and beech, growing on an acidic loam and on a calcareous sand, were exposed to increased CO₂ that was depleted in ¹³C. We traced the new C inputs from trees into dissolved organic C, into water‐extractable organic C, and into several particle‐size fractions. In addition, we incubated the labelled soils for 1 year and measured the production of dissolved organic C and CO₂ from new and old soil C. In the soil solutions of the topsoil, the dissolved organic C contained only 5–10% new C from the trees. The δ¹³C values of dissolved organic C resembled those of C pools smaller than 50 µm, which strongly suggests that the major source of dissolved organic C was humified old C. Apparently, throughfall, fresh litter and roots made only minor contributions to dissolved organic C. Water‐extractable organic C contained significantly larger fractions of new C than did the natural dissolved organic C (25–30%). The δ¹³C values of the water‐extractable organic C were closely correlated with those of sand fractions, which consisted of little decomposed organic carbon. The different origin of dissolved and water‐extractable organic C was also reflected in a significantly larger molar UV absorptivity and a smaller natural ¹³C abundance of dissolved organic C. This implies that the sampling method strongly influences the characteristics and sources of dissolved organic C. Incubation of soils showed that new soil C was preferentially respired as CO₂ and only a small fraction of new C was leached as dissolved organic C. Our results suggest that dissolved organic C is produced during incomplete decomposition of recalcitrant native C in the soils, whereas easily degradable new components are rapidly consumed by microbes and thus make only a minor contribution to the dissolved C fraction.     

STUDIES ON THE DECOMPOSITION OF PLANT MATERIAL IN SOIL

The organic matter in soils containing decomposing ¹⁴C-labelled ryegrass was fractionated chemically. Earlier work on these soils had shown that they contained a small fraction, heavily labelled relative to the rest of the soil organic matter, that was mineralized when the partially sterilized soils were incubated. Reagents effective in extracting heavily labelled-C included cold o.in HC1, boiling saturated CaSO₄ solution, and o.in Ba(OH)₂, but neither these nor any other reagent tested could extract material as heavily labelled as that mineralized when partially sterilized soil was incubated. Reagents that extract heavily labelled-C are poor extractants for humified material and are not strongly hydrolytic: the more vigorous the hydrolysis the smaller the proportion of labelled-C in the hydrolysate. The amounts of labelled-C dissolved by Ba(OH)₂ from soils sampled after different periods in the field were directly proportional to the amounts of labelled-C mineralized by those soils when partially sterilized (by exposure to CHC1₃ vapour), inoculated and incubated. Balance sheets are presented for the distribution of labelled and unlabelled-C in fractions separated by hydrolysis with 6N HC1, by NaOH extraction, by neutral pyrophosphate extraction, and by oxidation with H₂O₂. The fraction remaining after hydrolysis with 6N HC1 was the most lightly labelled and had the widest C/N ratio. The percentage of labelled-C in the material dissolved by alkali or by pyrophosphate was little more than in the material not dissolved, despite the presence in the soil of fractions differing at least twenty-fold in intensity of labelling.     

The effect of maize straw placement on mineralization of C and N in soil particle size fractions

Fundamental knowledge about the complex processes during the decomposition, mineralization and transfer of residue organic matter in soils is essential to assess risks of changes in agricultural practices. In a double tracer (¹³C, ¹⁵N) experiment the effect of maize straw on the mineralization dynamics and on the distribution of maize-derived organic matter within particle size fractions was investigated. Maize straw (a C₄ plant) labelled with ¹⁵N was added to soils (13.2 g dry matter kg^–1 soil) which previously had grown only C₃ plants, establishing two treatments: (i) soil mixed with maize straw (mixed), and (ii) soil with maize straw applied on the surface (surface). Samples were incubated in the laboratory at 14°C for 365 days. The size fractions (> 200 μm, 200–63 μm, 63–2 μm, 2–0.1 μm and < 0.1 μm), obtained after low-energy sonication (0.2 kJ g^–1), were separated by a combination of wet-sieving and centrifuging. The mineralization of maize C was similar in the two treatments after one year. However, decomposition of maize particulate organic matter (predominantly in the fraction > 200 μm) was significantly greater in the mixed treatment, and more C derived from the maize was associated with silt- and clay-sized particles. A two-component model fitted to the data yielded a rapidly mineralizable C pool (about 20% of total C) and a slowly mineralizable pool (about 80%). Generally, the size of the rapidly mineralizable C pool was rather small because inorganic N was rapidly immobilized after the addition of maize. However, the different mean half-lives of the C pools (rapidly decomposable mixed 0.035 years, and surface-applied 0.085 years; slowly decomposable mixed 0.96 years, and surface-applied 1.7 years) showed that mineralization was delayed when the straw was left on the surface. This seems to be because there is little contact between the soil microflora and plant residues. Evidently, the organic matter is more decomposed and protected within soil inorganic compounds when mixed into the soil than when applied on the soil surface, despite similar rates of mineralization.     

Influence of the rhizosphere on microbial biomass and recently formed organic matter

We have aimed to quantify the effect of roots on the size of the soil microbial biomass, and their influence on the turnover of soil organic matter and on the extent of the rhizosphere. We sampled a sandy clay loam topsoil from two subplots with different treatment histories. One had a normal arable fertilization record, the other had received only inorganic nitrogen fertilizer but no phosphorus and potassium for 30 years. Glucose labelled with ¹⁴C was added to both samples which were then incubated for 4 weeks before the soil was packed in cylinders and planted with ryegrass. In both soils, microbial biomass at the root surface doubled during the first 8 days. At day 15, the microbial biomass had further increased in the fertile soil, and the rhizosphere effect had extended 2.5 mm into the fertile soil, but to only 1 mm in the infertile soil. The microbial ¹⁴C increased threefold near the roots in the fertile soil as a result of assimilation of previously formed microbial residues, but in the infertile soil there was no increase. There was a close relation between the increase in microbial ¹⁴C and a decrease in ¹⁴C soluble in 2 m KCl, indicating that the microbial residues were more weakly adsorbed in the fertile soil. We conclude that the increased microbial population living near the root surfaces re‐assimilated part of the ¹⁴C‐labelled microbial residues in the fertile soil. In the infertile soil, microbial residues resisted decomposition because they were more strongly sorbed on to soil surfaces.     

Growing crops and transformations of 14C-labelled soil organic matter

Transformations of native soil organic materials, previously labelled with ¹⁴C, were investigated in soil planted with maize or perennial ryegrass and in fallow controls. There was more ¹⁴C in cold water extracts from planted soils than from fallow controls—an effect apparently caused by suppression of processes that remove labelled organic materials from this fraction of the soil organic matter. Decomposition of the labelled organic matter to ¹⁴CO₂ was significantly less in the planted soils than in fallow controls.     

Seasonal effects on photoassimilated carbon-14 in the root system of blue grama and associated soil organic matter

Blue grama (Bouteloua gracilis (H.B.K.) Lag.) was grown at day (14 h) and night temperatures of 25° and 15°C, respectively, in a ¹⁴CO₂-atmosphere during the last 31 of the 55 days from germination to seed set (period 1). An air-tight seal separated the shoot and root spheres. This period was followed by 21 days of a 14°C day (10 h) and 38°C night regime, and 29 days of continuous ?5°C (period 2), and 26 days of the original temperature and light conditions (period 3). Distribution of the assimilated ¹⁴C at the end of period 1 was: roots 33%; root-derived organic matter in the soil 23%; and 22% was released as CO₂. The washed root mass to root-derived soil organic matter ratio of the labelled ¹⁴C was 60 to 40. A root mass decrease of 45% over the cool and frost period changed this ratio to 23 to 77. Polysaccharides and 0.1 n NaOH-extractable organic matter decreased while potential dehydrogenase activity and total organic P increased during this same period, thereby confirming field related observations. Measured dehydrogenase activity overwinter may have two different origins. As total C content of the soil did not increase under the conditions of the experiment, it was postulated that a portion of the observed increase in total C in the field overwinter was of inorganic rather than organic origin.     

Rapid incorporation of carbon from fresh residues into newly formed stable microaggregates within earthworm casts

Earthworms play an important role in protecting carbon in the soil, but the exact influence of their activity on the distribution and protection of C is still poorly understood. We investigated the effect of earthworms on the formation of stable microaggregates inside newly formed macroaggregates and the distribution of C in them. We crushed (< 250 µm) soil, and subjected it to three treatments: (i) soil + ¹³C‐labelled residue + earthworms (these added after 8 days' incubation), (ii) soil + ¹³C‐labelled residue, and (iii) control (no additions), and then incubated it for 20 days. At the end, we measured the aggregate size distribution, total C and ¹³C, and we isolated microaggregates (53–250 µm) from macroaggregates (> 250 µm) formed. The ¹³C in fine particulate organic matter between and within the microaggregates was determined. Earthworms helped to form large macroaggregates (> 2000 µm). These large macroaggregates contained four times more stable microaggregates than those from samples without earthworms. There was more particulate organic matter within and between microaggregates in macroaggregates in the presence of earthworms. The larger amounts of organic matter inside stable microaggregates in casts than in bulk soil after 12 days of incubation (140 mg ¹³C kg^?1 soil compared with 20 mg ¹³C kg^?1 soil) indicates that these microaggregates are formed rapidly around freshly incorporated residues within casts. In conclusion, earthworms have a direct impact on the formation of stable microaggregates and the incorporation of organic matter inside these microaggregates, and it seems likely that their activity is of great significance for the long‐term stabilization of organic matter in soils.     

Simulating decomposition of 14C‐labelled fresh organic matter in bulk soil and soil particle fractions at various temperatures and moisture contents

¹⁴C‐labelled fresh organic matter (FOM) was homogeneously incorporated into an agricultural topsoil of small total organic carbon (TOC) content in order to perform decomposition batch experiments at temperatures (T) ranging from 5 to 45°C and soil gravimetric water contents (w) ranging from 7 to 35%. After 4–6‐month incubation (t_end), the residual ¹⁴C (D_end) was measured in bulk soil (0–2000 µm) and soil particle size fractions of 0–53, 53–200 and 200–2000 µm by chemical dispersion and sieving. The ¹⁴C‐FOM decomposition kinetics from soil were fitted either by a single first‐order reaction (rate constant, k_0–2000) assuming only a one‐pool model in the bulk soil or by consecutive first‐order reactions (rate constants, k_0–53 and k_53–2000) assuming a two‐pool model in the bulk soil aggregate structure. In the latter case, a two‐step reaction mechanism involving a FOM particle‐size decrease along the soil fractions was considered where k_0–53 was assumed to be a limiting rate constant. The ¹⁴C‐FOM decomposition kinetics was described for the experimental temperature and water ranges by Arrhenius and Michaelis‐Menten relationships, respectively. Additionally, the results obtained by the adapted Arrhenius physicochemical relationship were compared with the function proposed by Kirschbaum (1995) . Scaling functions T_m and w_m were established and can be used to simulate FOM decomposition rates under different temperature and moisture level conditions. Modelling based on consecutive first‐order reactions supported the hypothesis that the circulation (inflow and outflow) of C into the soil particle small‐size fractions (<53 µm) controls the total C mineralization.     

Seasonal transfers of assimilated 14C in grassland: Plant production and turnover,soil and plant respiration

Six areas of native grassland were labelled with ¹⁴C during a growing season. Transfers from the foliage to the roots and root respiration were measured. Plant production and turnover rates were determined by sampling the labelled material at different periods following exposure to ¹⁴CO₂.Above to beneath ground plant production ratios ranged between 1.1 and 1.9 with maximal translocation to the roots occurring during the drier summer months. The distribution of the photosynthates in the roots at different depths changed with time and soil moisture content. The upper part of the soil (0–10 cm) contained 49–77% of the labelled C found beneath the soil surface. Measurement of transfers with time of the above ground labelled C from living to dead plant and litter categories gave an insight into foliage dynamics and made it possible to estimate the seasonal shoot production at 130g Cm^?2 (1300kg ha^?1). Root growth represented 100g Cm^?2 (1000 kg ha^?1).Calculations of root and soil respiration were based on the CO₂ profiles in the soil. The fluxes of labelled and unlabelled CO₂ at the soil surface were estimated using the diffusion equation method. Respiration by roots and closely associated soil organisms accounted for 12 per cent of the net assimilation of CO₂ by the plants. This proportion was constant throughout the season and represented 19 per cent of the total CO₂ evolved at the soil surface.