Silver nanoparticles (AgNPs) are the most common nanomaterials incorporated in commercial products due to their antimicrobial activity. Recently, AgNPs were detected in surface waters suggesting the potential for bioavailability in the aquatic receptor organisms. This dissertation research attempts to understand the potential toxicity of AgNPs on water quality indicators, focusing on the microbial community and amphipods. This study evaluated whether: (1) the antimicrobial properties of AgNPs pose potential risks to microbial communities in pathogen impaired streams; (2) AgNPs can cause a shift in functional diversity and metabolic fingerprinting of microbial communities; (3) survival and growth of Hyalella azteca (amphipods) could be affected by AgNPs; and (4) surface coating agents influence AgNP toxicity in H. azteca. Microbial community responses to AgNPs were assessed using standard plate count, microbial enzyme assays, and carbon substrate utilization with Biolog EcoPlates™. Ten-day and 28-d toxicity tests were conducted in a static system to assess AgNP effects on H. azteca. AgNPs caused a 69% decrease in microbial concentration and a 77% decrease in β-glucosidase activity at 0.32 mg Ag kg-1 dry sediment. The substrate utilization pattern of the microbial community was altered by AgNPs at 0.33 mg Ag kg-1 dry sediment. Ten-day LC50s for the survival of H. azteca were 3.3, 9.2, and 230.0 µg Ag L-1 for AgNO3, citrate-AgNP, and PVP-AgNP, respectively, whereas the 28-d LC50s were 3.0, 3.5, and 66.0 µg Ag L-1 for AgNO3, citrate-AgNP, and PVP-AgNP, respectively. The EC20s for growth (calculated as biomass) for the 10-d test were 1.6, 4.7, and 188.1 µg Ag L-1 for AgNO3, citrate-AgNP, and PVP-AgNPs; while the 28-d EC20s for AgNO3, citrate-AgNP and PVP-AgNP were 3.2, 0.5, and < 50 µg Ag L-1. The NOECS for dry weight were 4 and 1, and 100 µg Ag L-1, while those for biomass were 2, 0.5, and < 50 µg Ag L-1 for AgNO3, citrate-AgNP, and PVP-AgNP, respectively. The overall toxicity followed the trend: AgNO3 > citrate-AgNP > PVP-AgNP. The studies suggest that AgNPs pose potential risks to microbial communities and epibenthic macroinvertebrates used as bioindicators of water quality to protect public health and ecosystem health.

京都大学 / 0048 / 新制・課程博士 / 博士(地球環境学) / 甲第17933号 / 地環博第112号 / 新制||地環||22(附属図書館) / 30753 / 京都大学大学院地球環境学舎環境マネジメント専攻 / (主査)教授 勝見 武, 教授 高岡 昌輝, 准教授 乾 徹 / 学位規則第4条第1項該当 / Doctor of Global Environmental Studies / Kyoto University / DFAM

[pt] A contaminação de solos por hidrocarbonetos é uma real preocupação ambiental em muitas partes do mundo devido a crescente dependência econômica dos derivados do petróleo, principalmente os combustíveis fósseis que estão frequentemente sujeitos a vazamentos e derramamentos acidentais. Estudos capazes de entender os mecanismos de biodegradação dos componentes da gasolina no solo não saturado se fazem necessários, pois auxiliam no processo de tomada de decisões em relação ao gerenciamento e controle da propagação em subsuperfície. No Brasil, o etanol é utilizado como aditivo oxigenado à gasolina e alguns autores sugerem para solo saturado que, por ser mais degradável, ele atrasa a degradação dos outros componentes mais tóxicos desse combustível. O trabalho anterior realizado por esse grupo de pesquisa para solo não saturado obteve conclusões semelhantes às já constatadas para solo saturado. No entanto, os mecanismos de degradação na zona não saturada ainda foram pouco estudados e compreendidos. Este estudo tem como objetivo analisar a degradação do contaminante em blocos não saturados indeformados de solo arenoso e siltoso, oriundos do município de Duque de Caxias – RJ, submetidos a um pulso de contaminante. Dois blocos, um arenoso (BA) e outro argiloso (BS), foram contaminados pela solução de dois porcento de Benzeno, quatro porcento de Tolueno em Heptano (BT); outros dois blocos, também um arenoso (EA) e outro argiloso (ES), foram submetidos à mesma solução adicionados etanol a vinte porcento (BTE). Foram realizados ensaios com a finalidade de monitorar a atividade degradadora total do meio, carbono disponível, perfil metabólico da microbiota, concentração dos contaminantes, além de medições da umidade volumétrica através do uso do Time Domain Reflectometer (TDR). A atividade microbiana inicial, antes da contaminação, apresentou valores baixos. Logo após a contaminação dos blocos, àqueles sujeitos a etanol apresentaram ausência de atividade enquanto os sujeitos apenas à mistura BT mostraram um aumento, contrariando as expectativas. As atividades microbianas oscilaram ao longo de todo o experimento o que pode indicar uma adaptação da microbiota às novas condições do meio. Com base nos resultados dos ensaios e monitoramentos realizados, podemos sugerir que o principal fator determinante para alteração da atividade foi à composição do solo. O contaminante não exerceu a influência esperada e vista nos estudos anteriores. / [en] The soil contamination by hydrocarbons is a real environmental concern in many parts of the world due to growing economic dependence on petroleum, mostly fossil fuels are often subject to leaks and accidental spills. Studies able to understand the mechanisms of biodegradation of gasoline components in unsaturated soil are necessary because they help in making decisions regarding the management and control of the propagation process in the subsurface. In Brazil, ethanol is used as an oxygenate gasoline additive and some authors suggest that for saturated soils, being more degradable, it delays the degradation of other more toxic components of this fuel. The previous study by this research group to unsaturated soil obtained similar results to those already observed for saturated soil. However, the mechanism of degradation in the unsaturated zone have few studies. This study aims to analyze the degradation mechanisms of the contaminant in unsaturated blocks of sand and silt soil from the city of Duque de Caxias - RJ, subjected to a pulse of contaminant. Two blocks, one sandy (BA) and silty (BS), were contaminated by two percent solution of benzene, four percent toluene in heptane (BT); other two blocks, also sandy (EA) and silty (ES), underwent the same solution added to twenty percent ethanol (BTE). Analyzes were performed to monitor the overall activity of the degrading medium, available carbon, the microbiota metabolic profile, concentration of the contaminants were performed as well as measurements of the water content through the use of Time Domain Reflectometer (TDR). The initial microbial activity before the contamination, showed low values. Soon after contamination of the blocks, those subjected to ethanol showed no activity while subject to BT mixture showed an increase, contrary to expectations. Microbial activities ranged throughout the experiment which may indicate an adaptation of microbes to new environmental conditions. Based on the results of testing and monitoring conducted, we suggest that the main determinant for changing the activity factor was the composition of the soil. The contaminant did not exert the expected influence and seen in previous studies

No description available.

A loss of soil organic matter (SOM), whether through natural means or management practices, results in soil degradation. Biochar as a soil amendment can alter soil properties, ultimately affecting the availability of nitrogen and water to plants and thus crop growth. The effects of biochar are not definitive, and often dependent on both the soil type and the biochar applied. Biochar properties can change according to the feedstock and production parameters, thus for their effective use further investigation is required to link biochar properties to its effects in soil. A high-temperature (600˚ C) biochar from a mixed-hardwood feedstock was investigated. The biochar increased the soil water retention, as demonstrated by a water release curve and field trials. This retention was predominant at higher water potentials, which was attributed to the greater number of meso (storage) pores in the biochar. Biochar did not affect the soil’s saturated hydraulic conductivity; this is thought to be due to the low number of macro (transmission) pores in the biochar. Thus there was no effect on the transmission rate in the soil. Biochar reduced gross ammonium levels in the soil via adsorption, but resulted in increased non-exchangeable ammonium levels, possibly due to physical entrapment. Where carbon was already abundant in the organically managed soil, the adsorbed ammonium reduced nitrification through lower substrate availability. The range of carbon fractions added as a result of the biochar amendment increased the total organic carbon (TOC) content of the soil, but this supplementary carbon was released by the microorganisms as carbon dioxide. Microorganisms in the relatively carbon poor conventionally managed soil (with lower TOC), assimilated the additional labile carbon increasing microbial biomass. The higher microbial biomass, combined with improvements in pH and the higher ammonium levels (as a result of the ammoniacal fertiliser) increased nitrification. These changes in water and nitrogen availability did not alter crop yields as measured in the glasshouse and field trials. The effects of this biochar in a sandy agricultural soil depended on the type and level of carbon and nitrogen present in the soil, thus consideration of these factors should be taken when applying.

Des études récentes ont permis de constater qu'une part de plus en plus importante des sols agricoles présente des déficiences en micronutriments. Ces carences posent donc de graves problèmes en terme de productivité. Il existe de nombreuses études sur le rôle des micronutriments dans le métabolisme des plantes et des microorganismes. Cependant aucune étude basée sur une approche holistique visant à prendre en considération l'ensemble des trois composantes biologiques du sol (plante, faune, microorganismes) par rapport à l'apport de micronutriments n'a été entreprise à ce jour. L'objectif de cette étude était de préciser l'impact des micronutriments sur les trois composantes biologiques d'un sol en étudiant différents paramètres tels que la production de biomasse végétale, les communautés bactériennes et les aspects fonctionnels à travers l'analyse de l'activité des enzymes du sol.Une expérimentation menée en champ a été réalisée sur une culture de colza. Une comparaison entre blocs parcellaires traités et non traité a été réalisé en analysant le nombre et le poids de siliques ; la teneur en chlorophylle des feuilles, le poids et la densité des vers de terre, les activités enzymatiques, la structure des communautés microbiennes et l'activité microbienne globale. L'apport de micronutriments induit des changements de tous les paramètres biologiques étudiés en fonction du stade de développement de la plante. Parmi les enzymes étudiées (phosphatases alcaline et acide, beta-glucosidase, alpha-glucosidase, beta xylosidase), une augmentation significative de la phosphatase alcaline et de l'alpha-glucosidase a été mesurée mais seulement lorsque la plante est au stade de grenaison. A ce stade, on constate également une augmentation de l'activité microbienne totale ainsi qu'une plus forte densité de vers de terre dans les blocs parcellaires traités. De plus, l'apport de micronutriments affecte de manière significative les communautés de bactéries nitrifiantes mais ce phénomène ne s'observe qu'au stade de la floraison. L'ensemble des ces résultats suggère donc que les micronutriments agissent comme des catalyseurs de l'activité biologique des différents acteurs du sol (plante, microorganismes et macrofaune). Ils sont à l'origine d'une succession d'inter action complexes et modifient alors les processus biologiques qui s'opèrent dans le sol.Le deuxième objectif de cette étude était de comparer l'impact de micronutriments sur les communautés microbiennes en présence /absence de plantes ou de vers de terre. Une expérimentation conduite en microcosme a été entreprise. Compte- tenu que les paramètres physico –chimiques du sol influent la biodisponibilité des micronutriments, l'expérience a été réalisée avec trois types de sols (sols limoneux, argileux et sableux). Les résultats obtenus mettent en évidence que l'impacte des micronutriments sur les communautés bactériennes est subordonné à la présence et le type d'organismes (vers de terre/plante) ainsi qu'à la nature du sol.Finalement, les micronutriments ne doivent plus être perçus en terme de productivité à travers une simple relation avec la plante mais au contraire comme des agents agissant comme catalyseur de l'activité biologique des principaux acteurs du sol. Une telle vision holistique constitue les premières données pour établir les bases de l'agroécologie. / Numerous arable lands in the world have a low availability of micronutrients. This lack of micronutrients has a serious impact on crop production. To increase plant productivity more attention must be paid to micronutrients. A number of studies have focused on the effect of micronutrients on plant and bacterial metabolism. However, studies on the impact of these micronutrients, when all living organisms (plant, macrofauna and microorganism) present in a field are taken into account, are scarce..The aim of the present study was to investigate the effects of these micronutrients on the three soil components. Different parameters were chosen such as plant productivity, bacterial community, soil enzyme and fauna to determine the impact of micronutrients on these organisms.A field experimental approach consisted of micronutrient treatments during oilseed rape cultivation in the Aisne Department (France) was carried out. In comparison to control plots, the weight and number of siliques (seed pods), the chlorophyll content of leaves, the weight and density of earthworms, soil enzyme activities, bacterial communities and total microbial activity were analysed. The addition of micronutrients caused changes to all the biological parameters studied in relation to the plant growth stage. Among the enzymes tested (alkaline and acid phosphatase, beta-glucosidase, alpha-glucosidase, beta xylosidase), a significant increase in alkaline phosphatase and alpha-glucosidase was measured at the 3rd stage of oilseed rape development with the addition of micronutrients. At this stage of oilseed rape development, total microbial activity and the weight and density of earthworms were higher in plots under micronutrient treatment. On the other hand, micronutrients had a significant impact on the nitrifying bacterial community and the total bacterial community only at the 2nd stage of oilseed rape development. All these results suggest that micronutrients act as a catalyst by stimulating plants, soil fauna and microorganisms directly or indirectly. Thus micronutrients can modify biological processes in all living organisms through successive complex interactions.The second objective of this study was to compare the impact of micronutrients on bacterial communities in presence/absence of plants or earthworms. An in vitro experimental approach using microcosms has been conducted to study different parameters (bacterial community, soil enzyme activity, total microbial activity). Because characteristics of soil contents are important in determining the availability of micronutrients in soils the experiment was conducted on three type of soil: sandy; loamy and clay soil. Our results bring to light that the application of micronutrients affected the bacterial communities. However the nature and the importance of this impact varies according to the organisms presents (plant or earthworms) and the nature of the soil.The results of this study provide a scientific basis for fertilizing and utilizing soil, protecting microbial diversity, and accomplishing the sustainable development of agroecology.

Les essences de forêts tropicales sont caractérisées par une importante variabilité de la qualité et de la stœchiométrie des feuilles qui tombent au sol. Les microorganismes hétérotrophes à la base des réseaux trophiques de décomposeurs dépendent principalement de ces ressources organiques qui varient de façon substantielle à petite échelle quant à la quantité et la contribution relative de certains éléments clés tels que le carbone (C), l'azote (N) et le phosphore (P). J'ai évalué au cours de cette thèse comment les variations de qualité et de stœchiométrie C:N:P de la ressource influençaient l'activité, la biomasse, la stœchiométrie et la structure des communautés des décomposeurs microbiens. J'ai réalisé ce travail en forêt Amazonienne de Guyane française sur des sols extrêmement appauvris en nutriments où les microorganismes hétérotrophes sont supposés être particulièrement dépendants du C et des nutriments provenant des litières. J'ai d'abord démontré que la qualité du C et le contenu en P des feuilles de litières expliquaient plus de 50% de la variabilité observée du processus de respiration microbien (SIR) du sol sous-jacent. Lors d'une expérience de fertilisation factorielle avec du C (sous forme de cellulose), de l'N (sous forme d'urée) et du P (sous forme de phosphate) sur le terrain, j'ai ensuite confirmé que la SIR de la communauté du sol était co-limitée par C et P, alors la SIR dans la litière était co-limitée par N et P. Ces limitations différentielles dans les litières et le sol sous-jacent étaient reliées à des modifications de la structure des communautés microbiennes, et en particulier des changements du ratio champignon:bactérie et de la proportion de bactéries copiotrophes et oligotrophes. Finalement au cours d'une expérience d'incubation au laboratoire, j'ai montré que la biomasse, la stœchiométrie et la structure des communautés microbiennes de la litière différaient fortement entre six litières chimiquement contrastées variant dans leur stœchiométrie initiale C:N:P. Cependant, les variations des paramètres microbiens étaient mieux expliqués par les caractéristiques de la fraction soluble des litières (y compris sa stœchiométrie) que par la qualité de la litière dans son ensemble, entrainant des variations de la stœchiométrie de la biomasse microbienne et un shift vers une dominance fongique en réponse à une augmentation de la stœchiométrie C:N:P des lessivâts. Collectivement, ces résultats montrent que des qualités de litière distinctes produites par une importante diversité d'essences forestières contrôlent la structure, la stœchiométrie, l'abondance et l'activité des communautés microbiennes des litières à petites échelles spatiales en forêt tropicale d'Amazonie. Par ailleurs, les litières en décomposition stimulent également les communautés microbiennes du sol sous-jacent, qui apparaissent être limitées par l'accès combiné à une source de C (énergie) et de P. L'importance de la contrainte stœchiométrique pour les microorganismes hétérotrophes à la base des réseaux trophiques de décomposeurs suggère que des modifications de la composition des communautés végétales ou des dépositions atmosphériques de N et/ou P peuvent avoir des conséquences plus lointaines sur les cycles du C et des nutriments au sein des biomes tropicaux. / Tree species-rich tropical rainforests are characterized by a high variability in quality and stoichiometry of leaf litter input to the soil. Microbial heterotrophs in the decomposer food web depend primarily on these organic resources that can vary dramatically in quantity, quality and relative contribution in key elements such as carbon (C), nitrogen (N), and phosphorus (P). I evaluated during this thesis how differences in leaf litter resource quality and C:N:P stoichiometry influence the activity, biomass, stoichiometry and community structure of microbial decomposers. I did this work in the Amazonian rainforest of French Guiana, where the soils are highly nutrient-impoverished and microbial heterotrophs are assumed to be particularly dependent on litter-derived nutrients. I first showed that leaf litter C quality and P content explained more than 50% of the observed variability of the microbial respiration process in the underlying soil. Using a fertilization experiment with C (as cellulose), N (as urea), and P (as phosphate) in the field, I further showed that microbial respiration process in the litter layer was co-limited by N and P, while that in the soil was co-limited by C and P. Additionally, distinct nutritional limitations in litter and underlying soil were related to shifts in the microbial community structure, especially regarding the fungi:bacteria ratio and the proportion of copiotrophic versus oligotrophic bacteria. Finally, during a laboratory incubation experiment, I showed that litter microbial biomass, stoichiometry and community structure differed strongly among leaf litter from six different tree species varying in C:N:P stoichiometry. The variations in microbial parameters among substrate litters, however, were not related to bulk leaf litter quality, but rather driven by the stoichiometry of the soluble fraction, with larger microbial C:nutrients ratios and a shift towards fungal dominance with increasing litter leachate C:N:P stoichiometry. Collectively, these results showed that the distinct leaf litter quality produced by a diverse tree canopy controls the structure, stoichiometry, abundance and activity of microbial communities in the studied Amazonian rainforest at small spatial scales. Moreover, the decomposing leaf litter stimulates microbial communities in the underlying soil that appear to be under the combined control of energy (C) and P availability. The strong stoichiometric constraint on microbial heterotrophs in the decomposer food web suggests far-ranging consequences on ecosystem C and nutrient cycling with ongoing alteration of nutrient deposition and tree species diversity in tropical rainforests..

La préservation de la qualité de l’eau est un enjeu majeur du 21ème siècle notamment avec l’augmentation de la population humaine. Or les rejets de stations d’épuration représentent un risque de pollution du milieu récepteur, notamment en azote et phosphore, aux effets néfastes pour la santé et l’environnement. C’est pour limiter cette pollution que les zones tampon végétalisées (ZTV), systèmes d’infiltration ou d’écoulement des effluents, plantés d’espèces végétales, ont été développées. Cette étude a pour objectif principal de mieux comprendre les transferts de nutriments dans ce type de système. L’impact des différents compartiments –Sol, végétation, microflore- a été étudié dans deux ZTVs de type « noues » situées en aval de stations d’épuration végétalisées en activité et dans une zone expérimentale aux modalités contrôlées. Cette étude a démontré l’importance de l’hydraulique du site, de la texture du sol et de la surface de la zone dans l’efficacité de rétention des flux d’eau et nutriments. L’impact du compartiment microbien dépend en grande partie du temps de rétention hydraulique lié à la surface de la ZTV, et du taux d’oxygénation de l’effluent. La végétation permet une amélioration de l’activité microbienne mais son rôle dans la rétention et l’élimination des nutriments s’avère mineur au regard des quantités en nutriments apportés. Le sol est le compartiment majeur de rétention du phosphore, même si une saturation rapide du sol en phosphore pourrait limiter cette potentialité. Ces observations ont permis de mettre en évidence une variabilité de l’efficacité des ZTVs dans la rétention de l’eau et des nutriments. Cette variabilité peut d’ores et déjà être minimisée par des mesures d’aménagement et de gestion. L’étude des transferts de nutriments mérite d’être approfondie et étendue à tous les types de ZTV. / The conservation of water quality is a major issue in the 21th century in particular with the increase in the human population. Water-treatment plants rejections represent a risk of pollution of the receiving environment, in particular in nitrogen and phosphorus, with fatal effects for the health and the environment. It is to limit this pollution that the vegetated buffer zones (VBZs), systems of infiltration, where water and nutrients retention are expected, were developed. This study has for main objective to better understand the transfers of nutrients in this type of system. The impact of the various compartments - soil, vegetation, microflora- was studied in two VBZs ditches type, situated downstream to vegetated water-treatment plants in service and in an experimental zone with controlled modalities. This study demonstrated the importance of site hydraulic, soil texture and VBZ surface in effectiveness of water and nutrients retention. The impact of microbial compartment depends largely on the hydraulic retention time bonds to VBZ surface and oxygenation rate of the effluent. Vegetation allows microbial activity improvement but its role in nutrients retention and removal is minor because of high quantities brought by effluents. The soil is the major compartment in phosphorus retention, although a rapid saturation of soil phosphorus could limit this potential. These observations demonstrate variability in the effectiveness of VBZ in water and nutrients retention. This variability could be minimized by planning and management measures. The study of nutrients transfers deserve to be further study and extended to all types of VBZ.

In der vorliegenden Arbeit wurden Untersuchungen zur mikrobiellen Aktivität von Pseudomonas putida DSM12735 durchgeführt. Als Messgröße diente die mikrobielle Wärmeleistung, basierend auf dem Stoffumsatz durch die Mikroorganismen. Ziel war es, die Vor- und Nachteile der verwendeten Kalorimeter herauszuarbeiten. Dafür wurden klassische Batch-Wachstumskurven aufgenommen. Ein weiteres Ziel bestand darin, eine Methode zur schnellen kalorimetrischen Detektion der mikrobiellen Aktivität insbesondere für die stationäre Phase zu entwickeln. In dieser Phase findet kein signifikanter Stoffumsatz statt. Durch das gezielte Auslösen einer zweiten Wachstumsphase und damit einem Stoffumsatz wird die mikrobielle Aktivität kalorimetrisch wieder messbar. Eingesetzt wurden folgende Kalorimeter: der Thermal Activity Monitor 2277 (TAM) mit den Kalorimetern Micro Reaction System 2250-4 ml und 2250-20 ml (kurz: TAM-4ml, TAM-20ml), das IC-Chip-Kalorimeter FCC22 (Institut für Physikalische Chemie, TU Freiberg) und das Kalorimeter Micro-DSC II (MDSC).

Currently, 74% of cotton acres in the United States are planted with glyphosatetolerant varieties. The average glyphosate-tolerant cotton crop is treated with glyphosate 2.1 times each year in addition to other herbicides, insecticides, and fungicides. The primary objectives of this research were to: 1) describe the influence of glyphosate and pesticides commonly applied at or near the time of cotton planting on soil microbial activity and biomass; 2) study the effect of glyphosate on fluometuron degradation; 3) evaluate the response of Rhizoctonia solani to glyphosate and fluometuron; 4) study changes in glyphosate metabolism that occur as a result of repeated glyphosate applications; and 5) define shifts in the soil microbial community. Additionally, methods for accelerated solvent extraction (ASE) of fluometuron from soils were developed. In one experiment, the addition of glyphosate reduced C-mineralization in soils treated with fluometuron, aldicarb, or mefenoxam + PCNB formulations. However, in a second experiment, C-mineralization increased when glyphosate was applied with fluometuron relative to fluometuron applied alone. Accelerated solvent extraction was used in experiments which demonstrated that application of glyphosate with fluometuron increased the rate of fluometuron degradation in soil relative to fluometuron alone. When glyphosate was added to minimal medium, degradation of fluometuron by R. solani was reduced and less fungal biomass was produced. The total amount of 14C-glyphosate mineralized was reduced when glyphosate was applied 5 times relative to 1, 2, 3, or 4 times. Incorporation of 14Cglyphosate residues into soil microbial biomass was greater following five glyphosate applications than one application 3 and 7 days after application (DAA). Soil fatty acid methyl ester (FAME) profiles were altered by five glyphosate applications relative to one application. Additionally, FAMEs common to gram-negative bacteria were present in higher concentrations following five applications relative to 1, 2, 3, or 4 applications both 7 and 14 DAA. These studies indicated that: 1) glyphosate altered the soil microbial response to other pesticides; 2) fluometuron-degrading microorganisms in soil responded differently to glyphosate; 3) changes in the dissipation or distribution of glyphosate following repeated glyphosate applications were associated with changes in the structural diversity of the soil microbial community.