Enhanced degradation of metalaxyl in Gley Humic and Dark Red Latosol

Enhanced degradation of the fungicide metalaxyl was investigated in two soils: a gley humic (GH) and a Dark Red Latosol (LE), collected at sites never exposed to the fungicide. The soil samples were treated with successive applications of metalaxyl as a commercial formulation and 14C-metalaxyl in laboratory. Metalaxyl biodegradation was analyzed during 63 days by means of radiometric techniques to verify biomineralization and degradation product formation from the applied 14C-metalaxyl. Although biomineralization (maximum of 14 and 8% in the GH and LE soils, respectively), and partial degradation (about 32 and 48%, respectively) were detected in both soils, enhanced degradation was verified only in the GH soil. Results proved that metalaxyl behaves differently in soils.

Modelling the risk of nitrate leaching from two soils amended with five different biosolids

High N concentrations in biosolids are one of the strongest reasons for their agricultural use. However, it is essential to understand the fate of N in soils treated with biosolids for both plant nutrition and managing the environmental risk of NO3--N leaching. This work aimed at evaluating the risk of NO3--N leaching from a Spodosol and an Oxisol, each one treated with 0.5-8.0 dry Mg ha-1 of fresh tertiary sewage sludge, composted biosolids, limed biosolids, heat-dried biosolids and solar-irradiated biosolids. Results indicated that under similar application rates NO3--N accumulated up to three times more in the 20 cm topsoil of the Oxisol than the Spodosol. However, a higher water content held at field capacity in the Oxisol compensated for the greater nitrate concentrations. A 20 % NO3--N loss from the root zone in the amended Oxisol could be expected. Depending on the biosolids type, 42 to 76 % of the NO3--N accumulated in the Spodosol could be expected to leach down from the amended 20 cm topsoil. NO3--N expected to leach from the Spodosol ranged from 0.8 (composted sludge) to 3.5 times (limed sludge) the amounts leaching from the Oxisol treated alike. Nevertheless, the risk of NO3--N groundwater contamination as a result of a single biosolids land application at 0.5-8.0 dry Mg ha-1 could be considered low.

Protective effect of divalent cations against aluminum toxicity in soybean

A large proportion of soybean fields in Brazil are currently cultivated in the Cerrado region, where the area planted with this crop is growing considerably every year. Soybean cultivation in acid soils is also increasing worldwide. Since the levels of toxic aluminum (Al) in these acid soils is usually high it is important to understand how cations can reduce Al rhizotoxicity in soybean. In the present study we evaluated the ameliorative effect of nine divalent cations (Ca, Mg, Mn, Sr, Sn, Cu, Zn, Co and Ba) in solution culture on Al rhizotoxicity in soybean. The growth benefit of Ca and Mg to plants in an acid Inceptisol was also evaluated. In this experiment soil exchangeable Ca:Mg ratios were adjusted to reach 10 and 60 % base saturation, controlled by different amounts of CaCl2 or MgCl2 (at proportions from 100:0 up to 0:100), without altering the soil pH level. The low (10 %) and adequate (60 %) base saturation were used to examine how plant roots respond to Al at distinct (Ca + Mg)/Al ratios, as if they were growing in soils with distinct acidity levels. Negative and positive control treatments consisted of absence (under native soil or undisturbed conditions) or presence of lime (CaCO3) to reach 10 and 60 % base saturation, respectively. It was observed that in the absence of Aluminum, Cu, Zn, Co and Sn were toxic even at a low concentration (25 µmol L-1), while the effect of Mn, Ba, Sr and Mg was positive or absent on soybean root elongation when used in concentrations up to 100 µmol L-1. At a level of 10 µmol L-1 Al, root growth was only reverted to the level of control plants by the Mg treatment. Higher Tin doses led to a small alleviation of Al rhizotoxicity, while the other cations reduced root growth or had no effect. This is an indication that the Mg effect is ion-specific and not associated to an electrostatic protection mechanism only, since all ions were divalent and used at low concentrations. An increased exchangeable Ca:Mg ratio (at constant soil pH) in the acid soil almost doubled the soybean shoot and root dry matter even though treatments did not modify soil pH and exchangeable Al3+. This indicates a more efficient alleviation of Al toxicity by Mg2+ than by Ca2+. The reason for the positive response to Mg2+ was not the supply of a deficient nutrient because CaCO3 increased soybean growth by increasing soil pH without inducing Mg2+ deficiency. Both in hydroponics and acid soil, the reduction in Al toxicity was accompanied by a lower Al accumulation in plant tissue, suggesting a competitive cation absorption and/or exclusion of Al from plant tissue stimulated by an Mg-induced physiological mechanism.

Simultaneous multielement extraction with the Mehlich-1 solution for Southern Brazilian soils determined by ICP-OES and the effects on the nutrients recommendations to crops

The amounts of macro (P, K, Ca and Mg) and micronutrients (Cu and Zn) extracted with the Mehlich-1 (M1) solution, by the 1.0 mol L-1 KCl (KCl) and with the 0.1 mol L-1 HCl (HCl) for representative soil types of the Rio Grande do Sul state (Brazil) were compared with those extracted with the Mehlich-1 solution determined with the inductively coupled plasma optical emission spectroscopy (ICP). The amounts of nutrients extracted by the different methods showed high correlation coefficients. On average, the Mehlich-1 solution extracted similar amounts of P, determined with colorimetric and ICP methods, and, K determined with emission and ICP. The amounts of Ca and Mg extracted with the Mehlich-1 solution, determined by ICP, were similar to those extracted with the KCl solution determined by the atomic absorption spectrophotometry. The amounts of Cu and Zn extracted with the Mehlich-1 solution, determined by the ICP, were higher than those extracted with the 0.1 mol L-1 HCl determined by the atomic absorption spectrophotometry. The results indicate that the Mehlich-1 solution and ICP can be used for simultaneous multielement extraction and determination for Southern Brazilian soils. However, a conversion factor for values interpretation is needed. The use of the conversion factor to determine the K availability index in soils is adequate and does not affect the K recommendations for crops in southern Brazilian soils.

In-furrow inoculation of soybean as alternative to fungicide and micronutrient seed treatment

Soybean is a major grain crop in Brazil, and yields can be considerably improved by inoculation with selected Bradyrhizobium strains. However, the incompatibility between inoculation and seed treatments with fungicides and micronutrients represents a major barrier to the achievement of high rates of biological N2 fixation. Inoculation practices that can alleviate the negative effects of agrochemicals must therefore be found and in-furrow inoculation seems to be an attractive alternative. This study reports the results of seven field experiments conducted in three growing seasons in Brazil; three in soils previously cropped with inoculated soybean (> 10(4) cells g-1 of soil of Bradyrhizobium), and four in areas where the crop was sown for the first time (< 10² cells g-1 of soil of Bradyrhizobium). The compatibility with fungicides and micronutrients was compared in seeds inoculated with peat or liquid inoculants, or treated with different doses of liquid inoculant in-furrow. In areas with established Bradyrhizobium populations, seed-applied agrochemicals did generally not affect nodulation, but also did not increase yields, while inoculation always increased N grain accumulation or yield, and N fertilizer decreased both nodulation and yield. Where soybean was sown for the first time, the seed treatment with agrochemicals affected nodulation when applied together with peat or liquid inoculant. In-furrow inoculation alleviated the effects of seed treatment with agrochemicals; the best performance was achieved with high Bradyrhizobium cell concentrations, with up to 2.5 million cells seed-1.

Phosphatase activity in sandy soil influenced by mycorrhizal and non-mycorrhizal cover crops

Cover crops may difffer in the way they affect rhizosphere microbiota nutrient dynamics. The purpose of this study was to evaluate the effect of mycorrhizal and non-mycorrhizal cover crops on soil phosphatase activity and its persistence in subsequent crops. A three-year experiment was carried out with a Typic Quartzipsamment. Treatments were winter species, either mycorrhizal black oat (Avena strigosa Schreb) or the non-mycorrhizal species oilseed radish (Raphanus sativus L. var. oleiferus Metzg) and corn spurry (Spergula arvensis L.). The control treatment consisted of resident vegetation (fallow in the winter season). In the summer, a mixture of pearl millet (Pennisetum americanum L.) with sunnhemp (Crotalaria juncea L.) or with soybean (Glycine max L.) was sown in all plots. Soil cores (0-10 cm) and root samples were collected in six growing seasons (winter and summer of each year). Microbial biomass P was determined by the fumigation-extraction method and phosphatase activity using p-nitrophenyl-phosphate as enzyme substrate. During the flowering stage of the winter cover crops, acid phosphatase activity was 30-35 % higher in soils with the non-mycorrhizal species oilseed radish, than in the control plots, regardless of the amount of P immobilized in microbial biomass. The values of enzyme activity were intermediate in the plots with corn spurry and black oat. Alkaline phosphatase activity was 10-fold lower and less sensitive to the treatments, despite the significant relationship between the two phosphatase activities. The effect of plant species on the soil enzyme profile continued in the subsequent periods, during the growth of mycorrhizal summer crops, after completion of the life cycle of the cover crops.

Geostatistical determination of heavy-metal distribution in a contaminated site - A case study

A two stage sampling strategy is necessary in order to optimize the study of distribution of pollution in soils and groundwater. First, detailed sampling from a limited area coupled with statistical analysis of the data are used to determine the microvariability of the parameter(s). The results from this detailed analysis are then used to calculate the optimal spacing between samples for the larger scale study. This two stage sampling strategy can result in significant financial savings during subsequent soil or groundwater remediation. This combined sampling and statistical analysis approach is illustrated with an example from a heavy metal contaminated site.

Distribution and vertical stratification of carbon and nitrogen in soil under different managements in the pampean region of Argentina

One of the expected benefits of no-tillage systems is a higher rate of soil C sequestration. However, higher C retention in soil is not always apparent when no-tillage is applied, due e.g., to substantial differences in soil type and initial C content. The main purpose of this study was to evaluate the potential of no-tillage management to increase the stock of total organic C in soils of the Pampas region in Argentina. Forty crop fields under no-tillage and conventional tillage systems and seven undisturbed soils were sampled. Total organic C, total N, their fractions and stratification ratios and the C storage capacity of the soils under different managements were assessed in samples to a depth of 30 cm, in three layers (0-5, 5-15 and 15-30 cm). The differences between the C pools of the undisturbed and cultivated soils were significant (p < 0.05) and most pronounced in the top (0-5 cm) soil layer, with more active C near the soil surface (undisturbed > no-tillage > conventional tillage). Based on the stratification ratio of the labile C pool (0-5/5-15 cm), the untilled were separated from conventionally tilled areas. Much of the variation in potentially mineralizable C was explained by this active C fraction (R² = 0.61) and by total organic C (R² = 0.67). No-till soils did not accumulate more organic C than conventionally tilled soils in the 0-30 cm layer, but there was substantial stratification of total and active C pools at no till sites. If the C stratification ratio is really an indicator of soil quality, then the C storage potential of no-tillage would be greater than in conventional tillage, at least in the surface layers. Particulate organic C and potentially mineralizable C may be useful to evaluate variations in topsoil organic matter.

Assessment of micronutrient extractants from soils of Paraná, Brazil

In Brazil, plant-available micronutrients in the soil can be determined by several chemical extractants, the most common of which are dilute acid and chelating solutions. The purpose of this study was to assess the extractants 0.1 mol L-1 HCl, Mehlich-1, Mehlich-3 and DTPA for analysis of the micronutrients Cu, Zn, Fe, and Mn in soils from the state of Paraná. In samples from 12 soils (0-20 cm layer), wheat was planted (Triticum aestivum), grown for 42 days after emergence, and then bean (Phaseolus vulgaris) for 38 days. At the end of each planting period, the soil was sampled again. All extractants tested to assess the availability of Cu, Zn, Fe, and Mn correlated with each other. The efficiency of the extractants HCl, Mehlich-3 and DTPA in assessing plant-available Cu was similar, unlike Mehlich-1, which proved less efficient. The extractants HCl, Mehlich-1 and Mehlich-3 were less efficient in estimating plant-available Zn and Fe, and the most indicated extractant is DTPA. The efficiency of the extractants HCl, Mehlich-1, Mehlich-3 and DTPA in assessing plant-available Mn in soils from Paraná was similar.

Characterization and nutrient release from silicate rocks and influence on chemical changes in soil

The expansion of Brazilian agriculture has led to a heavy dependence on imported fertilizers to ensure the supply of the growing food demand. This fact has contributed to a growing interest in alternative nutrient sources, such as ground silicate rocks. It is necessary, however, to know the potential of nutrient release and changes these materials can cause in soils. The purpose of this study was to characterize six silicate rocks and evaluate their effects on the chemical properties of treated soil, assessed by chemical extractants after greenhouse incubation. The experimental design consisted of completely randomized plots, in a 3 x 6 factorial scheme, with four replications. The factors were potassium levels (0-control: without silicate rock application; 200; 400; 600 kg ha-1 of K2O), supplied as six silicate rock types (breccia, biotite schist, ultramafic rock, phlogopite schist and two types of mining waste). The chemical, physical and mineralogical properties of the alternative rock fertilizers were characterized. Treatments were applied to a dystrophic Red-Yellow Oxisol (Ferralsol), which was incubated for 100 days, at 70 % (w/w) moisture in 3.7 kg/pots. The soil was evaluated for pH; calcium and magnesium were extracted with KCl 1 mol L-1; potassium, phosphorus and sodium by Mehlich 1; nickel, copper and zinc with DTPA; and the saturation of the cation exchange capacity was calculated for aluminum, calcium, magnesium, potassium, and sodium, and overall base saturation. The alternative fertilizers affected soil chemical properties. Ultramafic rock and Chapada mining byproduct (CMB) were the silicate rocks that most influenced soil pH, while the mining byproduct (MB) led to high K levels. Zinc availability was highest in the treatments with mining byproduct and Cu in soil fertilized with Chapada and mining byproduct.

Sulfur in agriculture

Sulfur (S) deficiency in soils is becoming increasingly common in many areas of the world as a result of agronomic practices, high biomass exportation and reduced S emissions to the atmosphere. In this review, the incidence and commercial exploitation of S pools in nature are discussed, as well as the importance of S for plants and the organic and inorganic S forms in soil and their transformations, especially the process of microbiological oxidation of elemental sulfur (S0) as an alternative to the replenishment of S levels in the soil. The diversity of S0-oxidizing microorganisms in soils, in particular the genus Thiobacillus, and the biochemical mechanisms of S0 oxidation in bacteria were also addressed. Finally, the main methods to measure the S0 oxidation rate in soils and the variables that influence this process were revised.

Particulate soil organic carbon and stratification ratio increases in response to crop residue decomposition under no-till

In soils under no-tillage (NT), the continuous crop residue input to the surface layer leads to carbon (C) accumulation. This study evaluated a soil under NT in Ponta Grossa (State of Paraná, Brazil) for: 1) the decomposition of black oat (Avena strigosa Schreb.) residues, 2) relation of the biomass decomposition effect with the soil organic carbon (SOC) content, the particulate organic carbon (POC) content, and the soil carbon stratification ratio (SR) of an Inceptisol. The assessments were based on seven samplings (t0 to t6) in a period of 160 days of three transects with six sampling points each. The oat dry biomass was 5.02 Mg ha-1 at t0, however, after 160 days, only 17.8 % of the initial dry biomass was left on the soil surface. The SOC in the 0-5 cm layer varied from 27.56 (t0) to 30.07 g dm-3 (t6). The SR increased from 1.33 to 1.43 in 160 days. There was also an increase in the POC pool in this period, from 8.1 to 10.7 Mg ha-1. The increase in SOC in the 0-5 cm layer in the 160 days was mainly due to the increase of POC derived from oat residue decomposition. The linear relationship between SOC and POC showed that 21 % of SOC was due to the more labile fraction. The results indicated that the continuous input of residues could be intensified to increase the C pool and sequestration in soils under NT.

EORTC 26083 phase I/II trial of dasatinib in combination with CCNU in patients with recurrent glioblastoma.

The treatment of patients with recurrent glioblastoma remains a major oncologic problem, with median survival after progression of 7-9 months. To determine the maximum tolerated dose and dose-limiting toxicity (DLT), the combination of dasatinib and cyclonexyl-chloroethyl-nitrosourea (CCNU) was investigated in this setting. The study was designed as multicenter, randomized phase II trial, preceded by a lead-in safety phase. The safety component reported here, which also investigated pharmacokinetics and preliminary clinical activity, required expansion and is therefore considered a phase I part to establish a recommended dosing regimen of the combination of CCNU (90-110 mg/m(2)) and dasatinib (100-200 mg daily). Overall, 28 patients were screened, and 26 patients were enrolled. Five dose levels were explored. DLTs, mainly myelosuppression, occurred in 10 patients. Grade 3 or 4 neutropenia was recorded in 7 patients (26.9%) and thrombocytopenia in 11 patients (42.3%). No significant effect of CCNU coadministration on dasatinib pharmacokinetics was found. Median progression-free survival (PFS) was 1.35 months (95% confidence interval: 1.2-1.4) and 6-month PFS was 7.7%. In this phase I study of recurrent glioblastoma patients, the combination of CCNU and dasatinib showed significant hematological toxicities and led to suboptimal exposure to both agents.

Quantification of aluminium in soil of the Solimões Formation, Acre State, Brazil

The variety of soils in the State of Acre is wide and their chemical profiles are still not fully understood. The nature of the material of origin of these soils is indicated by the high aluminium (Al) content, commonly associated with high calcium (Ca) and magnesium (Mg) contents. The study objective was to use different methods to quantify Al in soils from toposequences formed from material of a sedimentary nature originating from the Solimões Formation, in Acre, Brazil. Trenches were opened at three distinct points in the landscape: shoulder, backslope and footslope positions. Soil samples were collected for physical, chemical, mineralogical analyses. The Al content was quantified using different methods. High Al contents were found in most of these horizons, associated with high Ca and Mg levels, representing the predominant cations in the sum of exchangeable bases. The mineralogy indicates that the soils are still in a low weathering phase, with the presence of significant quantities of 2:1 minerals. Similar Al contents were determined by the methods of NaOH titration, xylenol orange spectrometry and inductively coupled plasma optical emission spectrometry. However, no consistent data were obtained by the pyrocatechol violet method. Extraction with KCl overestimated the exchangeable Al content due to its ability to extract the non-exchangeable Al present in the smectite interlayers. It was observed that high Al contents are related to the instability of the hydroxyl-Al smectite interlayers.

The role of fungi in the precipitation of calcite : relationships between fungal filaments, nanofibres, and needle fibre calcite

RésuméLes champignons sont impliqués dans les cycles biogéochimiques de différentes manières. En particulier, ils sont reconnus en tant qu'acteurs clés dans la dégradation de la matière organique, comme fournisseurs d'éléments nutritifs via l'altération des minéraux mais aussi comme grands producteurs d'acide oxalique et de complexes oxalo-métalliques. Toutefois, peu de choses sont connues quant à leur contribution à la genèse d'autres types de minéraux, tel que le carbonate de calcium (CaCO3). Le CaCO3 est un minéral ubiquiste dans de nombreux écosystèmes et il joue un rôle essentiel dans les cycles biogéochimiques du carbone (C) et du calcium (Ca). Le CaCO3 peut être d'origine physico-chimique ou biogénique et de nombreux organismes sont connus pour contrôler ou induire sa biominéralisation. Les champignons ont souvent été soupçonnés d'être impliqué dans ce processus, cependant il existe très peu d'informations pour étayer cette hypothèse.Cette thèse a eu pour but l'étude de cet aspect négligé de l'impact des champignons dans les cycles biogéochimiques, par l'exploration de leur implication potentielle dans la formation d'un type particulier de CaCO3 secondaires observés dans les sols et dans les grottes des environnements calcaires. Dans les grottes, ces dépôts sont appelés moonmilk, alors que dans les sols on les appelle calcite en aiguilles. Cependant ces deux descriptions correspondent en fait au même assemblage microscopique de deux habitus particulier de la calcite: la calcite en aiguilles (au sens strict du terme cette fois-ci) et les nanofibres. Ces deux éléments sont des habitus aciculaires de la calcite, mais présentent des dimensions différentes. Leur origine, physico-chimique ou biologique, est l'objet de débats intenses depuis plusieurs années déjà.L'observation d'échantillons environnementaux avec des techniques de microscopie (microscopie électronique et micromorphologie), ainsi que de la microanalyse EDX, ont démontré plusieurs relations intéressantes entre la calcite en aiguilles, les nanofibres et des éléments organiques. Premièrement, il est montré que les nanofibres peuvent être organiques ou minérales. Deuxièmement, la calcite en aiguilles et les nanofibres présentent de fortes analogies avec des structures hyphales, ce qui permet de confirmer l'hypothèse de leur origine fongique. En outre, des expériences en laboratoire ont confirmé l'origine fongique des nanofibres, par des digestions enzymatiques d'hyphes fongiques. En effet, des structures à base de nanofibres, similaires à celles observées dans des échantillons naturels, ont pu être produites par cette approche. Finalement, des enrichissements en calcium ont été mesurés dans les parois des hyphes et dans des inclusions intrahyphales provenant d'échantillons naturels de rhizomorphes. Ces résultats suggèrent une implication de la séquestration de calcium dans la formation de la calcite en aiguilles et/ou des nanofibres.Plusieurs aspects restent à élucider, en particulier la compréhension des processus physiologiques impliqués dans la nucléation de calcite dans les hyphes fongiques. Cependant, les résultats obtenus dans cette thèse ont permis de confirmer l'implication des champignons dans la formation de la calcite en aiguilles et des nanofibres. Ces découvertes sont d'une grande importance dans les cycles biogéochimiques puisqu'ils apportent de nouveaux éléments dans le cycle couplé C-Ca. Classiquement, les champignons sont considérés comme étant impliqués principalement dans la minéralisation de la matière organique et dans l'altération minérale. Cette étude démontre que les champignons doivent aussi être pris en compte en tant qu'agents majeurs de la genèse de minéraux, en particulier de CaCO3. Ceci représente une toute nouvelle perspective en géomycologie quant à la participation des champignons au cycle biologique du C. En effet, la présence de ces précipitations de CaCO3 secondaires représente un court-circuit dans le cycle biologique du C puisque du C inorganique du sol se retrouve piégé dans de la calcite plutôt que d'être retourné dans l'atmosphère.AbstractFungi are known to be involved in biogeochemical cycles in numerous ways. In particular, they are recognized as key players in organic matter recycling, as nutrient suppliers via mineral weathering, as well as large producers of oxalic acid and metal-oxalate. However, little is known about their contribution to the genesis of other types of minerals such as calcium carbonate (CaCO3). Yet, CaC03 are ubiquitous minerals in many ecosystems and play an essential role in the biogeochemical cycles of both carbon (C) and calcium (Ca). CaC03 may be physicochemical or biogenic in origin and numerous organisms have been recognized to control or induce calcite biomineralization. While fungi have often been suspected to be involved in this process, only scarce information support this hypothesis.This Ph.D. thesis aims at investigating this disregarded aspect of fungal impact on biogeochemical cycles by exploring their possible implication in the formation of a particular type of secondary CaC03 deposit ubiquitously observed in soils and caves from calcareous environments. In caves, these deposits are known as moonmilk, whereas in soils, they are known as Needle Fibre Calcite (NFC - sensu lato). However, they both correspond to the same microscopic assemblage of two distinct and unusual habits of calcite: NFC {sensu stricto) and nanofibres. Both features are acicular habits of calcite displaying different dimensions. Whether these habits are physicochemical or biogenic in origin has been under discussion for a long time.Observations of natural samples using microscopic techniques (electron microscopy and micromorphology) and EDX microanalyses have demonstrated several interesting relationships between NFC, nanofibres, and organic features. First, it has shown that nanofibres can be either organic or minera! in nature. Second, both nanofibres and NFC display strong structural analogies with fungal hyphal features, supporting their fungal origin. Furthermore, laboratory experiments have confirmed the fungal origin of nanofibres through an enzymatic digestion of fungal hyphae. Indeed, structures made of nanofibres with similar features as those observed in natural samples have been produced. Finally, calcium enrichments have been measured in both cell walls and intrahyphal inclusions of hyphae from rhizomorphs sampled in the natural environment. These results point out an involvement of calcium sequestration in nanofibres and/or NFC genesis.Several aspects need further investigation, in particular the understanding of the physiological processes involved in hyphal calcite nucleation. However, the results obtained during this study have allowed the confirmation of the implication of fungi in the formation of both NFC and nanofibres. These findings are of great importance regarding global biogeochemical cycles as they bring new insights into the coupled C and Ca cycles. Conventionally, fungi are considered to be involved in organic matter mineralization and mineral weathering. In this study, we demonstrate that they must also be considered as major agents in mineral genesis, in particular CaC03. This is a completely new perspective in geomycology regarding the role of fungi in the short-term (or biological) C cycle. Indeed, the presence of these secondary CaC03 precipitations represents a bypass in the short- term carbon cycle, as soil inorganic C is not readily returned to the atmosphere.