Le priming effect (PE) est la sur-minéralisation de la matière organique du sol (MOS) après un apport de matière organique fraiche. Ce phénomène serait généré par deux mécanismes distincts, la décomposition stœchiométrique et le « nutrient mining », ayant leur propre dynamique, leurs propres acteurs, leurs propres déterminants et leur propre stock de MO ciblés. Le premier serait plutôt lié à la séquestration de MO dans les sols et l’autre à sa déstabilisation. Comprendre comment piloter l’équilibre entre ces processus à travers les pratiques agricoles, permettrait d’améliorer durablement la fertilité des sols cultivés en milieu tropical dans un contexte de changement climatique. Pour identifier les déterminants, les acteurs et les effets du climat et des pratiques agricoles sur les différents processus générateurs de PE, nous avons combiné la caractérisation physicochimique des sols, la caractérisation des communautés microbiennes et le suivi de la minéralisation des MO par les techniques isotopiques. Ainsi, nous avons pu identifier différentes populations bactériennes et fongiques, associées à chacun des processus, que nous avons classées dans des guildes fonctionnelles. La taille de ces guildes déterminait l’équilibre entre les processus, et était corrélée à la qualité de la MO présente dans le sol. Plus précisément, nous avons montré que le PE stœchiométrique était favorisé dans les sols enrichis en matière organique peu évoluée et en nutriments N et P, entretenant donc une forte communauté de décomposeurs. Ces décomposeurs devaient ainsi limiter l’accès des mineurs à la nouvelle matière organique fraichement apportée et limiter le PE par « nutrient mining ». Sur parcelles agricoles, nos résultats suggéraient que le non labour, l’association légumineuses-céréales, et l’apport de compost favorisaient ces décomposeurs responsables du priming effect stœchiométrique et donc potentiellement la stabilisation durable de la matière organique dans les sols. / The priming effect (PE) is the supplementary mineralization of soil organic matter (MOS) after the addition of fresh organic matter. This phenomenon would be generated by two distinct mechanisms, stoichiometric decomposition and "nutrient mining", having their own dynamics, their own actors, their own determinants and their own MO stock targeted. The first would be related to the sequestration of MO in the soil and the other to its destabilization. Understanding how to manage the balance between these processes through agricultural practices, would allow to improve the fertility of soil cultivated in a tropical environment in a context of climate change.To identify the determinants, actors and effects of climate and agricultural practices on the various processes generating PE, we have combined soil physicochemical characterization, characterization of microbial communities and monitoring of mineralization of MO by isotopic techniques. Thus, we were able to identify different bacterial and fungal populations, associated with processes, which we classified in the functional guilds. The size of these guilds determined the balance between the processes, and was correlated with the quality of the MO present in the soil. Specifically, we showed that stoichiometric PE was favored in soils enriched with high quality organic matter and nutrients, N and P, thus maintaining a strong community of decomposers. These decomposers also limit the access of miners to the provided new organic matter hence limiting PE by "nutrient extraction". On agricultural plots, our results suggest that non-tillage, legume-cereal rotations and compost amendments favor these decomposers responsible for the stoichiometric priming effect and thus potentially the long-term stabilization of organic matter in soils.

Les objectifs de cette thèse étaient (i) d'étudier les mécanismes par lesquels les nématodes bactérivores régulent les flux de nutriments (N et P) dans la rhizosphère du riz pluvial dans un sol ferrallitique de Madagascar, (ii) d'étudier les effets des interactions bactéries – bactérivores sur les fonctions de la plante (croissance et nutrition), et (iii) d'évaluer les principaux facteurs contrôlant la boucle microbienne des sols dans une optique d'intensification. Notre modèle biologique comprenait Acrobeloides sp. (Cephalobidae) et Oryza sativa (Poaceae).Deux voies ont été proposées pour expliquer les effets positifs des bactérivores sur la disponibilité des nutriments et les fonctions de la plante: voie de minéralisation et voie d'exploration. Afin d'identifier l'implication de chaque voie dans les flux de P inorganique à l'interface sol-plante nous avons utilisé le radio-isotope 32P comme traceur. Lorsque le pH du sol est corrigé par l'ajout de dolomie, la présence des nématodes bactérivores améliore la minéralisation nette de P, la production de biomasse et la nutrition de la plante sans modifier la ramification du système racinaire, contrairement à ce qui a été observé par certains auteurs en présence de protistes. En effet, en présence de nématodes et de dolomie, la L-value dans les parties aériennes de la plante augmente de 49%, la biomasse totale de la plante augmente de 22%, et la quantité de P total dans les tissus de la plante augmente de 9%. Ainsi, Acrobeloides sp. stimule le prélèvement de P par la plante via la voie nutritionnelle ou voie de minéralisation.Les effets de Acrobeloides sp. sur les fonctions de la plante sont variables (positives, neutres, négatives), en fonction des pratiques agricoles et des caractéristiques abiotiques du sol. Tout d'abord, ces effets peuvent dépendre de la variété de riz puisque l'attractivité des nématodes Acrobeloides sp. par la rhizosphère du riz varie en fonction des variétés de riz pluvial, plus précisément en fonction de la composition et de la quantité d'exsudats racinaires libérés par la plante. Ensuite, l'activité mutualiste des nématodes est influencée (maximisée) par la présence d'arbres (agroforesterie) dans la parcelle de riz pluvial. En plus des plantes, les paramètres édaphiques étaient des facteurs clés. L'activité mutualiste des nématodes est fortement limitée par le pouvoir fixateur des Ferralsols. Ainsi, l'ajout de dolomie limite la capacité de sorption de P du sol et améliore l'activité mutualiste des nématodes bactérivores. Enfin, l'activité mutualiste des nématodes sur la croissance et la nutrition du riz pluvial est favorisée par les valeurs élevées de pH et de teneur en Mg dans le sol. Pour conclure nous avons trouvé qu'il est possible d'intensifier l'activité mutualiste des nématodes bactérivores dans un sol tropical pauvre à travers des pratiques agricoles spécifiques (gestion des sols, agroforesterie). / Using Acrobeloides sp. (Cephalobidae) and Oryza sativa (Poaceae), we aimed at (i) studying the effects of bacterivorous nematodes on plant functions growing in a nutrient-poor soil in Madagascar, (ii) testing the mechanisms involved in the mutualistic activity of the bacterivorous nematodes in a strong P-depleted Ferralsol from the highlands of Madagascar, and (iii) identifying the agronomic and edaphic factors that can drive the soil microbial loop.Two pathways have been proposed to explain the positive effects of bacterial-feeders on nutrients availability and plant functions: mineralization pathway and exploration pathway. In order to test the involvement of each pathway in the mutualistic activity of bacterivorous nematodes, we assessed the effect of nematodes inoculation on inorganic P flows from soil to plant using the 32P labelling technique. When the soil pH was corrected with dolomite lime, we showed that the presence of Acrobeloides sp. improved net P mineralization and plant (Oryza sativa) functions (growth and nutrition) but did not alter lateral root growth as protists do. Indeed, in the presence of nematodes and dolomite, the shoot L-value increased by 49%, the plant total biomass increased by 22% and the plant total P amount increased by 9%. Thus, Acrobeloides sp. increased plant P uptake through the "nutritional" or "mineralization" pathway.The effects of Acrobeloides sp. on Oryza sativa functions were variable (positive, neutral or negative) according to agricultural practices and soil abiotic variables. First, these effects may depend on the rice cultivar used in the experiment. Indeed, the nematodes attractancy to root-adhering soils varied according to upland rice cultivar, probably according to the amount and composition of root exudates. The mutualistic activity of nematodes is also influenced (maximized) by the presence of trees (agroforestry) in the rice field. Besides plants, the abiotic soil parameters were key drivers. The mutualistic activity of nematodes was strongly limited by the ability of Ferralsols to precipitate and chemisorb phosphates. The addition of dolomite lime limited phosphates sorption on soil and enhanced the mutualistic activity of the nematodes. Also, the high values of soil pH and Mg content tended to increase the mutualistic activity of the bacterivorous nematodes on rice growth and nutrition. To conclude, we found that it is possible to drive the mutualistic activity of bacterivorous nematodes in poor tropical soils through specific soil and agricultural practices.

Les Ferralsols se caractérisent par une microstructure très développée dont dépend étroitement leur fertilité. Cette microstructure repose sur des microagrégats très stables, de 50 à 800 µm de diamètre. Sous végétation naturelle comme sous pâturage, la densité apparente des sols, la taille et le mode d'assemblage des microagrégats, ainsi que la caractérisation de la matière organique, ont permis de mettre en évidence d'importantes variations avec la profondeur. Nous avons établi une typologie du mode d'assemblage des microagrégats ainsi qu'un modèle mettant en relation structure et porosité. La matière organique se singularise, en pyrolyse, par une composante de haute température, d'origine conjecturale, mais très probablement due à un effet de matrice minérale. L'ensemble des variations observées ne peuvent a priori s'expliquer par des différences d'usage du sol, mais plutôt grâce à l'activité des insectes sociaux qui sont vraisemblablement à l'origine des microagrégats.

Submitted by Biblioteca de Pós-Graduação em Geoquímica BGQ (bgq@ndc.uff.br) on 2016-04-29T16:09:29Z No. of bitstreams: 1 Tese_Ricardo Cesar-2014.pdf: 8808586 bytes, checksum: 7553914038f43221f7bee0f88ba00197 (MD5) / Made available in DSpace on 2016-04-29T16:09:29Z (GMT). No. of bitstreams: 1 Tese_Ricardo Cesar-2014.pdf: 8808586 bytes, checksum: 7553914038f43221f7bee0f88ba00197 (MD5) / Coordenação de Aperfeiçoamento de Pessoal de Nível Superior / Conselho Nacional de Desenvolvimento Científico e Tecnológico / Universidade Federal Fluminense. Instituto de Química. Programa de Pós-Graduação em Geociências-Geoquímica. Niterói, RJ / O presente trabalho trata da avaliação do risco ecológico associado à disposição de sedimentos de dragagem em latossolos e chernossolos, com base em indicadores pedogeoquímicos e ecotoxicológicos. Para tanto, uma amostra de sedimento dragado oriundo do canal do Cunha (Baía de Guanabara-RJ) foi coletada. Estudos preliminares foram conduzidos com uma amostra de lodo de esgoto, de forma a gerar dados-base que subdiassem a adoção de estratégias analíticas com o sedimento dragado. Além da avaliação dos patógenos, foi realizada a caracterização física, química e mineralógica dos materiais. Bioensaios agudos e crônicos com organismos terrestres (minhocas, colêmbolos, enquitreídeos e alface) e aquáticos (microcrustáceos e algas) foram realizados em misturas solo:resíduo. As doses de sedimento (expressas em %) capazes de causar efeitos significativos a 50% dos organismos-teste (CL50 ou CE50) foram estatisticamente estimadas. Os indicadores foram numericamente integrados visando a construção de um índice de risco ecotoxicológico. Os teores de zinco, cobre, chumbo, mercúrio e cromo no sedimento excederam as concentrações estipuladas pela legislação brasileira. As propriedades dos solos estudados desempenharam papel fundamental na redução dos níveis de ecotoxicidade, com especial atenção aos argilominerais 2:1 e aos altos níveis de fertilidade, capazes de atenuar a ecotoxicidade das misturas de Chernossolo. Com o lodo, a reprodução de algas foi o indicador mais sensível (CE50 = 1,61% em Latossolo). Nas misturas de solo:sedimento, a reprodução de algas, a germinação de alface e a imobilidade de microcrustáceos foram os indicadores mais sensíveis, com CE50 em torno de 3,5% em Latossolo. A estimativa numérica do risco revelou níveis maiores de ecotoxicidade para os tratamentos de Chernossolo. Quando os ensaios de reprodução com colêmbolos são excluídos da análise, os riscos são maiores em Latossolo. Tal fato pode estar associado ao fato de que os colêmbolos estão geralmente menos sensíveis à água intertiscial do solo, se comparados aos demais organismos-teste / This work aims to evaluate the environmental risks associated with the disposal of dredged sediments in ferralsols and chernosols, based on pedogeochemical and ecotoxicological indicators. A sample of dredged sediment was collected in the Cunha channel (Guanabara Bay-RJ). Preliminary studies were conducted with a sewage sludge sample (SL), in order to establish analytical estrategies to be adopted in the assays with the sediment. In addition to the pathogens, the physical, chemical and mineralogical properties of the materials were characterized. Acute and chronic bioassays with terrestrial (earthworms, springtails, enchytraeids and lettuce) and aquatic (micro-crustaceans and algae) organisms were performed in mixtures of residue:solo. The doses of sediment (in %) required to cause significant effects on 50% of the exposed organisms (LC50 or EC50) were statistically estimated. The indicators were numerically integrated to estimate the potential risks, based on the construction of an ecotoxicological risk index. The concentrations of zinc, copper, lead, mercury and chromium in the dredged sediment were higher than the limits established by Brazilian legislation. Soil properties played an important role in the reduction of ecotoxicity levels, especially 2:1 clay minerals and high levels of fertility, which were able to reduce the ecotoxicity levels in chernosol mixtures. The assays with SL revealed that the reprocuction of algae was the most sensible indicator (EC50 = 1.61% in ferralsol). In mixtures of soil:sediment, the reproduction of algae, germination of lettuce and immobility of microcrustaceans were the most sensible indicators (EC50 ~ 3.5% in ferralsol). The numerical estimation of the potential risks demonstrated the occurrence of higher levels of ecotoxicity for the chernosols treatments. When the reproduction assay with springtails was excluded from the analysis, the values of risk were higher for the ferralsol mixtures. Such fact is due to the lower sensibility of springtails to soil pore water, compared to the other testorganisms

Latossolos com horizonte A húmico (LH) são solos com importantes reservas de carbono orgânico (CO) em profundidade. Esses solos são comuns nas regiões Sudeste e Sul do Brasil, com menor ocorrência no Nordeste. Aspectos da gênese e preservação da matéria orgânica (MO) desses solos não foram completamente entendidos ainda. Particularmente interessantes são os processos responsáveis pelo espessamento (hiperdesenvolvimento) e estabilidade do CO do horizonte húmico. A hipótese de que a MO do horizonte húmico tenha importante contribuição de carbono (C) de origem pirogênica é testada. O objetivo deste trabalho foi, portanto, caracterizar quimicamente a MO do horizonte húmico e relacionar os resultados com a gênese e preservação do CO. Oito pedons de diferentes localidades do país (São Paulo, Rio de Janeiro, Minas Gerais, Paraná, Santa Catarina e Pernambuco) sob vegetação nativa foram empregados no estudo. Descrições morfológicas e análises de caracterização de rotina dos solos foram realizadas. Além disso, ácidos húmicos (AH) foram extraídos do horizonte húmico e analisados por ressonância magnética nuclear de 13C, enquanto que as frações leves (livre e oclusa), huminas e extratos alcalinos da MO dos mesmos pedons foram analisados através da pirólise. Os pedons possuem feições morfológicas conspícuas, como horizonte A hiperdesenvolvido, variando de 120 cm a 220 cm de espessura, e cores escuras em profundidade. Os horizontes húmicos são argilosos a muitos argilosos, fortemente ácidos e álicos. As formas de CO oxidável com dicromato respondem por 2/3 da reserva de carbono total (CT) do solo. Caulinita foi o principal mineral identificado por difratometria de raios X na fração argila desses indivíduos, seguido por gibbsita e, em alguns pedons, vermiculita com hidroxientrecamadas. Correlações entre atributos do solo e diferentes formas de C oxidável revelaram-se significativas apenas para alumínio pobremente cristalino extraído com oxalato de amônio ácido (r2 = 0,69), que se apresenta como um dos mecanismos de estabilização do C. Os AH apresentam estrutura química distinta entre os pedons e também variam com a profundidade, indicando, sobretudo o efeito do tipo de vegetação. Nos pedons estudados há maior concentração relativa de C alquila em superfície e C O-alquilas e C aromático em subsuperfície. A abundância de C O-alquilas sugere que boa parte da MO do horizonte húmico tem um grau de humificação pouco avançado. Os principais produtos da pirólise das frações leves, extratos alcalinos e huminas são carboidratos, compostos alifáticos e aromáticos, mas em proporções diferentes. As amostras da fração leve livre são mais ricas em resíduos frescos derivados de plantas, com presença de ligninas, enquanto que na fração leve oclusa (FLO) preponderam compostos aromáticos e alifáticos. Os extratos são enriquecidos em compostos de nitrogênio e carboidratos. A hipótese de uma MO altamente recalcitrante, formada predominantemente por produtos da decomposição de carvão foi rejeitada, já que produtos similares a esses foram encontrados apenas na FLO, que tem pouca importância no conteúdo de CT do solo. A estabilização da MO desses LH depende de 14 uma combinação de abundante entrada de biomassa e da atuação de mecanismos que retardam a mineralização da MO, como associações organominerais. / Ferralsols with umbric epipedons are soils with important organic carbon (OC) reserves in depth. These soils are common in the Southeast and South regions of Brazil, with fewer occurrences in the Northeast region. Both genesis and organic matter (OM) accumulation aspects of these soils were not completely understood yet. Particularly interesting are the soil processes responsible for thickening (over development) and for OC stability of the umbric horizon. The hypothesis that the OM from umbric epipedon has important pyrogenic carbon (C) contribution is tested. The aim of this work, therefore, was to characterize the chemistry of OM from umbric epipedon and to relate the results with soil genesis and OC preservation. Eigth pedons from different states of Brazil (São Paulo, Rio de Janeiro, Minas Gerais, Paraná, Santa Catarina and Pernambuco) under native vegetation were used in the study. Morphological descriptions and routine soil characterization analysis were carried out. Furthermore, humic acids (HA) were extracted from umbric horizon and analyzed by solid state 13C nuclear magnetic resonance, whereas light fractions (free and occluded), humin and alkaline extracts of the OM of the same pedons were analyzed by pyrolysis. The pedons have conspicuous morphological features, such as a developed umbric epipedon, ranging from 120 cm to 220 cm of thickness, and dark colors in depth. The umbric horizons are clayey to heavy clayey, strongly acidic and alic. Dichromate oxidable OC contents account for 2/3 of the total carbon (TC) soil reserve. Kaolinite is the main mineral identified by X ray diffractometry in the clay fraction, followed by gibbsite and vermiculite. Correlations between soil attributes and different forms of oxidable C were significant only for poorly crystalline aluminum forms extracted with oxalate ammonium acid (r2 = 0,69), which could be one of the C stabilization mechanisms. HA shown distinct chemical structures between pedons and also in depth, indicating the effect of the vegetation type. In the studied pedons there are higher relative concentration of alkyl C in surface and O-alkyl C and aromatic C in subsurface. Light fractions, alkaline extracts and humins main pyrolysis products are carbohydrates, aliphatic, and aromatic compounds, but in different proportions. In free light fraction (FLF) fresh residues derived from plants are common, with presence of lignin, whereas in the occluded light fraction (OLF), aromatic and aliphatic compounds predominate. Extracts are enriched in carbohydrates and N compounds. The hypothesis of a highly recalcitrant MO, produced by the fragments of charcoal in partial decomposition has been rejected, since they were found only into OLF, which has little importance in the TC content of soil. The stabilization of organic matter in these Ferralsols depends on a combination of abundant biomass input and the action of mechanisms to retard the mineralization of OM, such as a strong association with clay minerals.

Latossolos com horizonte A húmico (LH) são solos com importantes reservas de carbono orgânico (CO) em profundidade. Esses solos são comuns nas regiões Sudeste e Sul do Brasil, com menor ocorrência no Nordeste. Aspectos da gênese e preservação da matéria orgânica (MO) desses solos não foram completamente entendidos ainda. Particularmente interessantes são os processos responsáveis pelo espessamento (hiperdesenvolvimento) e estabilidade do CO do horizonte húmico. A hipótese de que a MO do horizonte húmico tenha importante contribuição de carbono (C) de origem pirogênica é testada. O objetivo deste trabalho foi, portanto, caracterizar quimicamente a MO do horizonte húmico e relacionar os resultados com a gênese e preservação do CO. Oito pedons de diferentes localidades do país (São Paulo, Rio de Janeiro, Minas Gerais, Paraná, Santa Catarina e Pernambuco) sob vegetação nativa foram empregados no estudo. Descrições morfológicas e análises de caracterização de rotina dos solos foram realizadas. Além disso, ácidos húmicos (AH) foram extraídos do horizonte húmico e analisados por ressonância magnética nuclear de 13C, enquanto que as frações leves (livre e oclusa), huminas e extratos alcalinos da MO dos mesmos pedons foram analisados através da pirólise. Os pedons possuem feições morfológicas conspícuas, como horizonte A hiperdesenvolvido, variando de 120 cm a 220 cm de espessura, e cores escuras em profundidade. Os horizontes húmicos são argilosos a muitos argilosos, fortemente ácidos e álicos. As formas de CO oxidável com dicromato respondem por 2/3 da reserva de carbono total (CT) do solo. Caulinita foi o principal mineral identificado por difratometria de raios X na fração argila desses indivíduos, seguido por gibbsita e, em alguns pedons, vermiculita com hidroxientrecamadas. Correlações entre atributos do solo e diferentes formas de C oxidável revelaram-se significativas apenas para alumínio pobremente cristalino extraído com oxalato de amônio ácido (r2 = 0,69), que se apresenta como um dos mecanismos de estabilização do C. Os AH apresentam estrutura química distinta entre os pedons e também variam com a profundidade, indicando, sobretudo o efeito do tipo de vegetação. Nos pedons estudados há maior concentração relativa de C alquila em superfície e C O-alquilas e C aromático em subsuperfície. A abundância de C O-alquilas sugere que boa parte da MO do horizonte húmico tem um grau de humificação pouco avançado. Os principais produtos da pirólise das frações leves, extratos alcalinos e huminas são carboidratos, compostos alifáticos e aromáticos, mas em proporções diferentes. As amostras da fração leve livre são mais ricas em resíduos frescos derivados de plantas, com presença de ligninas, enquanto que na fração leve oclusa (FLO) preponderam compostos aromáticos e alifáticos. Os extratos são enriquecidos em compostos de nitrogênio e carboidratos. A hipótese de uma MO altamente recalcitrante, formada predominantemente por produtos da decomposição de carvão foi rejeitada, já que produtos similares a esses foram encontrados apenas na FLO, que tem pouca importância no conteúdo de CT do solo. A estabilização da MO desses LH depende de 14 uma combinação de abundante entrada de biomassa e da atuação de mecanismos que retardam a mineralização da MO, como associações organominerais. / Ferralsols with umbric epipedons are soils with important organic carbon (OC) reserves in depth. These soils are common in the Southeast and South regions of Brazil, with fewer occurrences in the Northeast region. Both genesis and organic matter (OM) accumulation aspects of these soils were not completely understood yet. Particularly interesting are the soil processes responsible for thickening (over development) and for OC stability of the umbric horizon. The hypothesis that the OM from umbric epipedon has important pyrogenic carbon (C) contribution is tested. The aim of this work, therefore, was to characterize the chemistry of OM from umbric epipedon and to relate the results with soil genesis and OC preservation. Eigth pedons from different states of Brazil (São Paulo, Rio de Janeiro, Minas Gerais, Paraná, Santa Catarina and Pernambuco) under native vegetation were used in the study. Morphological descriptions and routine soil characterization analysis were carried out. Furthermore, humic acids (HA) were extracted from umbric horizon and analyzed by solid state 13C nuclear magnetic resonance, whereas light fractions (free and occluded), humin and alkaline extracts of the OM of the same pedons were analyzed by pyrolysis. The pedons have conspicuous morphological features, such as a developed umbric epipedon, ranging from 120 cm to 220 cm of thickness, and dark colors in depth. The umbric horizons are clayey to heavy clayey, strongly acidic and alic. Dichromate oxidable OC contents account for 2/3 of the total carbon (TC) soil reserve. Kaolinite is the main mineral identified by X ray diffractometry in the clay fraction, followed by gibbsite and vermiculite. Correlations between soil attributes and different forms of oxidable C were significant only for poorly crystalline aluminum forms extracted with oxalate ammonium acid (r2 = 0,69), which could be one of the C stabilization mechanisms. HA shown distinct chemical structures between pedons and also in depth, indicating the effect of the vegetation type. In the studied pedons there are higher relative concentration of alkyl C in surface and O-alkyl C and aromatic C in subsurface. Light fractions, alkaline extracts and humins main pyrolysis products are carbohydrates, aliphatic, and aromatic compounds, but in different proportions. In free light fraction (FLF) fresh residues derived from plants are common, with presence of lignin, whereas in the occluded light fraction (OLF), aromatic and aliphatic compounds predominate. Extracts are enriched in carbohydrates and N compounds. The hypothesis of a highly recalcitrant MO, produced by the fragments of charcoal in partial decomposition has been rejected, since they were found only into OLF, which has little importance in the TC content of soil. The stabilization of organic matter in these Ferralsols depends on a combination of abundant biomass input and the action of mechanisms to retard the mineralization of OM, such as a strong association with clay minerals.