Trading nitrogen for carbon: Nitrogen and carbon translocation in a plant/fungal (Metarhizium spp.) symbiosis

While nitrogen is critical for all plants, they are unable to utilize organically bound nitrogen in soils. Therefore, the majority of plants obtain useable nitrogen through nitrogen fixing bacteria and the microbial decomposition of organic matter. In the majority of cases, symbiotic microorganisms directly furnish plant roots with inorganic forms of nitrogen. More than 80% of all land plants form intimate symbiotic relationships with root colonizing fungi. These common plant/fungal interactions have been defined largely through nutrient exchange, where the plant receives limiting soil nutrients, such as nitrogen, in exchange for plant derived carbon. Fungal endophytes are common plant colonizers. A number of these fungal species have a dual life cycle, meaning that they are not solely plant colonizers, but also saprophytes, insect pathogens, or plant pathogens. By using 15N labeled, Metarhizium infected, wax moth larvae (Galleria mellonella) in soil microcosms, I demonstrated that the common endophytic, insect pathogenic fungi Metarhizium spp. are able to infect living soil borne insects, and subsequently colonize plant roots and furnish ts plant host with useable, insect-derived nitrogen. In addition, I showed that another ecologically important, endophytic, insect pathogenic fungi, Beauveria bassiana, is able to transfer insect-derived nitrogen to its plant host. I demonstrated that these relationships between various plant species and endophytic, insect pathogenic fungi help to improve overall plant health. By using 13C-labeled CO2, added to airtight plant growth chambers, coupled with nuclear magnetic resosnance spectroscopy, I was able to track the movement of carbon from the atmosphere, into the plant, and finally into the root colonized fungal biomass. This indicates that Metarhizium exists in a symbiotic partnership with plants, where insect nitrogen is exchanged for plant carbon. Overall these studies provide the first evidence of nutrient exchange between an insect pathogenic fungus and plants, a relationship that has potentially useful implications on plant primary production, soil health, and overall ecosystem stability.

Caractérisation du cycle et des sources d'azote dans les lacs tempérés par l'utilisation d'isotopes stables

Nous avons étudié l’application de plusieurs mesures d’isotopes stables afin de caractériser les processus du cycle de l’azote et les sources d’azote dans les lacs tempérés à diverses échelles spatiales et temporelles. Les résultats d’une étude à travers 65 lacs sur un gradient trophique ont démontré que le ratio d’isotopes stables d’azote (δ15N) des sédiments de surface est un indicateur de l’importance relative des sources d’azote anthropique, mais que ce ratio peut être altéré par la diagenèse. La mesure du δ15N des sédiments demeure néanmoins un outil permettant de déterminer à long terme le changement des charges en azote anthropique aux écosystèmes lacustres et les causes de l’eutrophisation de ces systèmes. Nos résultats d’une étude sur la variation saisonnière de plusieurs isotopes stables dans trois lacs peu profonds situés sur un gradient trophique et ayant différents régimes de stratification ont démontré que cette approche est prometteuse dans les lacs mésotrophes et stratifiés. Dans ces systèmes, le δ15N de la matière organique particulaire (MOP) aurait le potentiel de déterminer les sources d’azote assimilées par le phytoplancton. Cependant les mesures d’isotopes stables du carbone (δ13C) et du ratio C:N indiquent que les apports de matières organiques du bassin versant peuvent altérer les relations observées. Nous avons également constaté une déviation de la relation 1:1 entre les isotopes stables d’azote et d’oxygène (δ18O) du nitrate (NO3-) indiquant son assimilation et sa nitrification simultanée. Cette application est particulièrement prometteuse puisque la nitrification est méconnue dans les lacs et peut exacerber les effets de l’eutrophisation.

Mercure, arsenic et sélénium au Burkina Faso : bioaccumulation, transfert trophique dans les systèmes aquatiques et évaluation de bioaccessibilité chez les humains

L’extraction aurifère est l’une des activités humaines qui a fortement accru l’émission de contaminants métalliques dans l’environnement. Le mercure (Hg), l’arsenic (As) et le sélénium (Se) sont 3 polluants métalliques de grande toxicité environnementale. En milieu aquatique, ils peuvent subir des transformations menant à des composés capables de bioaccumulation et de bioamplification. Il peut en résulter des concentrations 106 fois celle mesurée dans l’eau chez les poissons et les organismes situés en haut des chaînes alimentaires posant de ce fait de graves menaces pour la santé de ces organismes ainsi que leurs consommateurs y compris les humains. Cette étude a évalué les teneurs en Hg, As et Se dans les milieux aquatiques au Burkina Faso, une région d’Afrique sub-saharienne soumise à une exploitation minière intensive. Le risque potentiel pour les organismes aquatiques et les humains a été évalué en considérant les effets des interactions antagonistes Se/Hg et As/Se. La bioaccumulation et le transfert du Hg et du Se dans les réseaux trophiques sont également décrits. L’exposition au Hg de poissons par les humains a été également évalué au laboratoire par mesure de la bioaccessibilité comme équivalent de la biodisponibilité par simulation de la digestion humaine. En général, les milieux aquatiques étudiés étaient peu affectés par ces 3 métal(loïd)s bien que certaines espèces de poisson issus des réservoirs les plus profonds indiquent des teneurs de Hg au dessus de 500 ngHg/g (poids frais) recommandé par l’OMS. Ces niveaux sont susceptibles de présenter des risques toxicologiques pour les poissons et pour leurs consommateurs. En considérant l’antagonisme Se/Hg, 99 % des échantillons de poisson seraient moins exposés à la toxicité du Hg dû à la présence simultanée du sélénium dans le milieu et pourraient être consommés sans risque. Cependant, les effets potentiels de l’antagonisme As/Se pourraient réduire les effets bénéfiques du Se et ramener cette proportion à 83 %. L’application des mesures de signatures en isotopes stables d’azote (δ15N) et de carbone (δ13C) des organismes aquatiques a permis le traçage des voies de transfert du Hg et du Se dans les réseaux trophiques. On y observe des chaînes trophiques très courtes (3 - 4 niveaux trophiques) et des poissons majoritairement benthiques. L’approche isotopique n’a cependant pas permis de détecter les variations saisonnières des niveaux de contamination en Hg des poissons. L’exploration des contenus stomacaux des poissons a permis de mieux expliquer la baisse des concentrations en Hg et Se observées chez certains poissons au cours de la saison sèche en lien avec la variation de la composition des proies que l’analyse isotopique n’a pas cerné. L’étude suggère que l’analyse de contenus stomacaux ainsi que l’étude de la dynamique des communautés d’invertébrés couplées à celle des métaux pourraient améliorer la compréhension du fonctionnement des écosystèmes étudiés. Enfin, l’évaluation expérimentale de l’exposition au Hg indique que les modes de traitement avant consommation ainsi que l’usage de composés alimentaires tels le thé, le café lors de repas de poisson par certaines communautés humaines ont un impact sur la bioaccessibilité du Hg de poisson. Ces résultats, sous réserve de validation par des modèles animaux, suggèrent la prise en compte des habitudes alimentaires des communautés dans l’élaboration adéquat des avis de consommation de poisson.

Essai de détection de processus migratoires à travers les isotopes de strontium et d’oxygène : étude des restes humains du cimetière Saint-Matthew (Québec, 1771-1860)

Afin de distinguer les immigrants de première génération des individus nés à Québec et de discuter de l’identité des immigrants de cette ville aux XVIIIe et XIXe siècles, trente-quatre squelettes humains exhumés du cimetière protestant Saint-Matthew (Québec, 1771-1860) ont fait l’objet d’analyses ostéologiques et isotopiques du strontium (87Sr/86Sr) et de l’oxygène (δ18O). Les teneurs obtenues, bien que moins précises que les données historiques, ont permis de distinguer trois groupes d’origine, soit les individus nés à Québec (N = 12), les immigrants de première génération provenant le plus probablement des îles Britanniques et du nord de la France (N = 19) et les immigrants de première génération dont l’origine ne peut être précisée (N = 3). De plus, l’origine écossaise de certains individus a pu être suggérée en fonction de compositions isotopiques variant entre -10,0 et -9,09 % vs VSMOW. La comparaison des groupes d’origine à des données provenant de sources historiques et d’une étude antérieure a permis de dresser un portrait de l’identité des immigrants, à la fois sur les plans populationnel et individuel. De plus, les compositions isotopiques (δ18O, 87Sr/86Sr, δ13C et δ15N) nous laissent croire qu’au moins un individu pourrait être d’origine amérindienne et qu’un autre proviendrait d’une partie de l’Europe plus appauvrie en 18O (possiblement un pays scandinave ou une région alpine). La distribution spatiale des sépultures nous a également permis d’émettre des hypothèses sur les liens familiaux et sociaux d’immigrants inhumés en caveaux ou entassés de façon particulièrement modeste.

Écologie et implications trophiques de la cyanobactérie Lyngbya wollei dans le fleuve Saint-Laurent

Les proliférations nuisibles de la cyanobactérie filamenteuse benthique Lyngbya wollei qui forme des tapis déposés sur les sédiments ont augmenté en fréquence au cours des 30 dernières années dans les rivières, lacs et sources de l'Amérique du Nord. Lyngbya wollei produit des neurotoxines et des composés organiques volatils (géosmin, 2-méthylisobornéol) qui ont des répercussions sur la santé publique de même que des impacts d'ordre socioéconomiques. Cette cyanobactérie est considérée comme un habitat et une source de nourriture de piètre qualité pour les invertébrés en raison de sa gaine robuste et de sa production de toxines. Les proliférations de L. wollei ont été observées pour la première fois en 2005 dans le fleuve Saint-Laurent (SLR; Québec, Canada). Nous avons jugé important de déterminer sa distribution sur un tronçon de 250 km afin d'élaborer des modèles prédictifs de sa présence et biomasse en se basant sur les caractéristiques chimiques et physiques de l'eau. Lyngbya wollei était généralement observé en aval de la confluence de petits tributaires qui irriguent des terres agricoles. L’écoulement d’eaux enrichies à travers la végétation submergée se traduisait par une diminution de la concentration d’azote inorganique dissous (DIN), alors que les concentrations de carbone organique dissous (DOC) et de phosphore total dissous (TDP) demeuraient élevées, produisant un faible rapport DIN :TDP. Selon nos modèles, DOC (effet positif), TP (effet négatif) et DIN :TDP (effet négatif) sont les variables les plus importantes pour expliquer la répartition de cette cyanobactérie. La probabilité que L. wollei soit présent dans le SLR a été prédite avec exactitude dans 72 % à 92 % des cas pour un ensemble de données indépendantes. Nous avons ensuite examiné si les conditions hydrodynamiques, c'est-à-dire le courant généré par les vagues et l'écoulement du fleuve, contrôlent les variations spatiales et temporelles de biomasse de L. wollei dans un grand système fluvial. Nous avons mesuré la biomasse de L. wollei ainsi que les variables chimiques, physiques et météorologiques durant trois ans à 10 sites le long d'un gradient d'exposition au courant et au vent dans un grand (148 km2) lac fluvial du SLR. L'exposition aux vagues et la vitesse du courant contrôlaient les variations de biomasses spatiales et temporelles. La biomasse augmentait de mai à novembre et persistait durant l'hiver. Les variations interannuelles étaient contrôlées par l'écoulement de la rivière (niveau d'eau) avec la crue printanière qui délogeait les tapis de l'année précédente. Les baisses du niveau d'eau et l'augmentation de l'intensité des tempêtes anticipées par les scénarios de changements climatiques pourraient accroître la superficie colonisée par L. wollei de même que son accumulation sur les berges. Par la suite, nous avons évalué l'importance relative de L. wollei par rapport aux macrophytes et aux épiphytes. Nous avons examiné l'influence structurante de l'échelle spatiale sur les variables environnementales et la biomasse de ces producteurs primaires (PP) benthiques. Nous avons testé si leur biomasse reflétait la nature des agrégats d'habitat basées sur l'écogéomorphologie ou plutôt le continuum fluvial. Pour répondre à ces deux questions, nous avons utilisé un design à 3 échelles spatiales dans le SLR: 1) le long d'un tronçon de 250 km, 2) entre les lacs fluviaux localisés dans ce tronçon, 3) à l'intérieur de chaque lac fluvial. Les facteurs environnementaux (conductivité et TP) et la structure spatiale expliquent 59% de la variation de biomasse des trois PP benthiques. Spécifiquement, les variations de biomasses étaient le mieux expliquées par la conductivité (+) pour les macrophytes, par le ratio DIN:TDP (+) et le coefficient d'extinction lumineuse (+) pour les épiphytes et par le DOC (+) et le NH4+ (-) pour L. wollei. La structure spatiale à l'intérieur des lacs fluviaux était la plus importante composante spatiale pour tous les PP benthiques, suggérant que les effets locaux tels que l'enrichissement par les tributaire plutôt que les gradients amont-aval déterminent la biomasse de PP benthiques. Donc, la dynamique des agrégats d'habitat représente un cadre général adéquat pour expliquer les variations spatiales et la grande variété de conditions environnementales supportant des organismes aquatiques dans les grands fleuves. Enfin, nous avons étudié le rôle écologique des tapis de L. wollei dans les écosystèmes aquatiques, en particulier comme source de nourriture et refuge pour l'amphipode Gammarus fasciatus. Nous avons offert aux amphipodes un choix entre des tapis de L. wollei et soit des chlorophytes filamenteuses ou un tapis artificiel de laine acrylique lors d'expériences en laboratoire. Nous avons aussi reconstitué la diète in situ des amphipodes à l'aide du mixing model (d13C et δ15N). Gammarus fasciatus choisissait le substrat offrant le meilleur refuge face à la lumière (Acrylique>Lyngbya=Rhizoclonium>Spirogyra). La présence de saxitoxines, la composition élémentaire des tissus et l'abondance des épiphytes n'ont eu aucun effet sur le choix de substrat. Lyngbya wollei et ses épiphytes constituaient 36 et 24 % de l'alimentation in situ de G. fasciatus alors que les chlorophytes, les macrophytes et les épiphytes associées représentaient une fraction moins importante de son alimentation. Les tapis de cyanobactéries benthiques devraient être considérés comme un bon refuge et une source de nourriture pour les petits invertébrés omnivores tels que les amphipodes.

High-momentum proton removal from 16 O and the (e,e'p) cross section

The cross section for the removal of high-momentum protons from 16O is calculated for high missing energies. The admixture of high-momentum nucleons in the 16O ground state is obtained by calculating the single-hole spectral function directly in the finite nucleus with the inclusion of short-range and tensor correlations induced by a realistic meson-exchange interaction. The presence of high-momentum nucleons in the transition to final states in 15N at 60¿100 MeV missing energy is converted to the coincidence cross section for the (e,e¿p) reaction by including the coupling to the electromagnetic probe and the final state interactions of the outgoing proton in the same way as in the standard analysis of the experimental data. Detectable cross sections for the removal of a single proton at these high missing energies are obtained which are considerably larger at higher missing momentum than the corresponding cross sections for the p-wave quasihole transitions. Cross sections for these quasihole transitions are compared with the most recent experimental data available.

Root effects on the turnover of grain legume residues in soil

Das Ziel dieser Arbeit war, die Einflüsse von Wurzeln und Rhizodeposition auf den Umsatz von Körnerleguminosenresiduen und damit verknüpfte mikrobielle Prozesse zu untersuchen. In einem integrierten Versuch wurden Ackerbohne (Vicia faba L.), Erbse (Pisum sativum L.) und Weiße Lupine (Lupinus albus L.) untersucht. Der Versuch bestand aus drei Teilen, zwei Gefäß-Experimenten und einem Inkubationsexperiment, in denen ausgehend von einem Gefäß-Experiment derselbe Boden und dasselbe Pflanzenmaterial verwendet wurden. In Experiment I wurde die Stickstoff-Rhizodeposition der Körnerleguminosenarten, definiert als wurzelbürtiger N nach dem Entfernen aller sichtbaren Wurzeln im Boden, gemessen und der Verbleib des Rhizodepositions-N in verschiednenen Bodenpools untersucht. Dazu wurden die Leguminosen in einem Gefäßversuch unter Verwendung einer in situ 15N-Docht-Methode mit einer 15N Harnstofflösung pulsmarkiert. In Experiment II wurde der Umsatz der N-Rhizodeposition der Körnerleguminosen und der Einfluss der Rhizodeposition auf den anschließenden C- und N-Umsatz der Körnerleguminosenresiduen in einem Inkubationsexperiment untersucht. In Experiment III wurde der N-Transfer aus den Körnerleguminosenresiduen einschließlich N-Rhizodeposition in die mikrobielle Biomasse und die Folgefrüchte Weizen (Triticum aestivum L.) und Raps (Brassica napus L.) in einem Gewächshaus-Gefäßversuch ermittelt. Die in situ 15N Docht-Markierungs-Methode wies hohe 15N Wiederfindungsraten von ungefähr 84 Prozent für alle drei Leguminosenarten auf und zeigte eine vergleichsweise homogene 15N Verteilung zwischen verschiedenen Pflanzenteilen zur Reife. Die Wurzeln zeigten deutliche Effekte auf die N-Dynamik nach dem Anbau von Körnerleguminosen. Die Effekte konnten auf die N-Rhizodeposition und deren anschließenden Umsatz, Einflüsse der Rhizodeposition von Körnerleguminosen auf den anschließenden Umsatz ihrer Residuen (Stängel, Blätter, erfassbare Wurzeln) und die Wirkungen nachfolgender Nichtleguminosen auf den Umsatzprozess der Residuen zurückgeführt werden: Die N-Rhizodeposition betrug zur Reife der Pflanzen bezogen auf die Gesamt-N- Aufnahme 13 Prozent bei Ackerbohne und Erbse und 16 Prozent bei Weißer Lupine. Bezogen auf den Residual N nach Ernte der Körner erhöhte sich der relative Anteil auf 35 - 44 Prozent. Die N-Rhizodeposition ist daher ein wesentlicher Pool für die N-Bilanz von Körnerleguminosen und trägt wesentlich zur Erklärung positiver Fruchtfolgeeffekte nach Körnerleguminosen bei. 7 - 21 Prozent des Rhizodepositions-N wurden als Feinwurzeln nach Nasssiebung (200 µm) wiedergefunden. Nur 14 - 18 Prozent des Rhizodepositions-N wurde in der mikrobiellen Biomasse und ein sehr kleiner Anteil von 3 - 7 Prozent in der mineralischen N Fraktion gefunden. 48 bis 72 Prozent der N-Rhizodeposition konnte in keinem der untersuchten Pools nachgewiesen werden. Dieser Teil dürfte als mikrobielle Residualmasse immobilisiert worden sein. Nach 168 Tagen Inkubation wurden 21 bis 27 Prozent des Rhizodepositions-N in den mineralisiert. Der mineralisierte N stammte im wesentlichen aus zwei Pools: Zwischen 30 Prozent und 55 Prozent wurde aus der mikrobiellen Residualmasse mineralisiert und eine kleinere Menge stammte aus der mikrobielle Biomasse. Der Einfluss der Rhizodeposition auf den Umsatz der Residuen war indifferent. Durch Rhizodeposition wurde die C Mineralisierung der Leguminosenresiduen nur in der Lupinenvariante erhöht, wobei der mikrobielle N und die Bildung von mikrobieller Residualmasse aus den Leguminosenresiduen in allen Varianten durch Rhizodepositionseinflüsse erhöht waren. Das Potential des residualen Körnerleguminosen-N für die N Ernährung von Folgefrüchten war gering. Nur 8 - 12 Prozent des residualen N wurden in den Folgenfrüchten Weizen und Raps wiedergefunden. Durch die Berücksichtigung des Rhizodepositions-N war der relative Anteil des Residual-N bezogen auf die Gesamt-N-Aufnahme der Folgefrucht hoch und betrug zwischen 18 und 46 Prozent. Dies lässt auf einen höheren N-Beitrag der Körnerleguminosen schließen als bisher angenommen wurde. Die residuale N-Aufnahme von Weizen von der Blüte bis zur Reife wurde durch den Residual-N gespeist, der zur Blüte in der mikrobiellen Biomasse immobilisiert worden war. Die gesamte Poolgröße, Residual-N in der mikrobiellen Biomasse und in Weizen, veränderte sich von der Blüte bis zur Reife nicht. Jedoch konnte ein Rest von 80 Prozent des Residual-N in keinem der untersuchten Pools nachgewiesen werden und dürfte als mikrobielle Residualmasse immobilisiert worden sein oder ist noch nicht abgebaut worden. Die zwei unterschiedlichen Folgefrüchte - Weizen und Raps - zeigten sehr ähnliche Muster bei der N-Aufnahme, der Residual-N Wiederfindung und bei mikrobiellen Parametern für die Residuen der drei Körnerleguminosenarten. Ein differenzierender Effekt auf den Umsatz der Residuen bzw. auf das Residual-N-Aneignungsvermögen der Folgefrüchte konnte nicht beobachtet werden.

Composition and stability of organic matter fractions in soils under different land use regimes

Soil organic matter (SOM) vitally impacts all soil functions and plays a key role in the global carbon (C) cycle. More than 70% of the terrestric C stocks that participate in the active C cycle are stored in the soil. Therefore, quantitative knowledge of the rates of C incorporation into SOM fractions of different residence time is crucial to understand and predict the sequestration and stabilization of soil organic carbon (SOC). Consequently, there is a need of fractionation procedures that are capable of isolating functionally SOM fractions, i.e. fractions that are defined by their stability. The literature generally refers to three main mechanisms of SOM stabilization: protection of SOM from decomposition by (i) its structural composition, i.e. recalcitrance, (ii) spatial inaccessibility and/or (iii) interaction with soil minerals and metal ions. One of the difficulties in developing fractionation procedures for the isolation of functional SOM fractions is the marked heterogeneity of the soil environment with its various stabilization mechanisms – often several mechanisms operating simultaneously – in soils and soil horizons of different texture and mineralogy. The overall objective of the present thesis was to evaluate present fractionation techniques and to get a better understanding of the factors of SOM sequestration and stabilization. The first part of this study is attended to the structural composition of SOM. Using 13C cross-polarization magic-angle spinning (CPMAS) nuclear magnetic resonance (NMR) spectroscopy, (i) the effect of land use on SOM composition was investigated and (ii) examined whether SOM composition contributes to the different stability of SOM in density and aggregate fractions. The second part of the present work deals with the mineral-associated SOM fraction. The aim was (iii) to evaluate the suitability of chemical fractionation procedures used in the literature for the isolation of stable SOM pools (stepwise hydrolysis, treatments using oxidizing agents like Na2S2O8, H2O2, and NaOCl as well as demineralization of the residue obtained by the NaOCl treatment using HF (NaOCl+HF)) by pool sizes, 13C and 14C data. Further, (iv) the isolated SOM fractions were compared to the inert organic matter (IOM) pool obtained for the investigated soils using the Rothamsted Carbon Model and isotope data in order to see whether the tested chemical fractionation methods produce SOM fractions capable to represent this pool. Besides chemical fractionation, (v) the suitability of thermal oxidation at different temperatures for obtaining stable SOC pools was evaluated. Finally, (vi) the short-term aggregate dynamics and the factors that impact macroaggregate formation and C stabilization were investigated by means of an incubation study using treatments with and without application of 15N labeled maize straw of different degradability (leaves and coarse roots). All treatments were conducted with and without the addition of fungicide. Two study sites with different soil properties and land managements were chosen for these investigations. The first one, located at Rotthalmünster, is a Stagnic Luvisol (silty loam) under different land use regimes. The Ah horizons of a spruce forest and continuous grassland and the Ap and E horizons of two plots with arable crops (continuous maize and wheat cropping) were examined. The soil of the second study site, located at Halle, is a Haplic Phaeozem (loamy sand) where the Ap horizons of two plots with arable crops (continuous maize and rye cropping) were investigated. Both study sites had a C3-/C4-vegetational change on the maize plot for the purpose of tracing the incorporation of the younger, maize-derived C into different SOM fractions and the calculation of apparent C turnover times of these. The Halle site is located near a train station and industrial areas, which caused a contamination with high amounts of fossil C. The investigation of aggregate and density fractions by 13C CPMAS NMR spectroscopy revealed that density fractionation isolated SOM fractions of different composition. The consumption of a considerable part (10–20%) of the easily available O-alkyl-C and the selective preservation of the more recalcitrant alkyl-C when passing from litter to the different particulate organic matter (POM) fractions suggest that density fractionation was able to isolate SOM fractions with different degrees of decomposition. The spectra of the aggregate fractions resembled those of the mineral-associated SOM fraction obtained by density fractionation and no considerable differences were observed between aggregate size classes. Comparison of plant litter, density and aggregate size fractions from soil under different land use showed that the type of land use markedly influenced the composition of SOM. While SOM of the acid forest soil was characterized by a large content (> 50%) of POM, which contained high amounts of spruce-litter derived alkyl-C, the organic matter in the biologically more active grassland and arable soils was dominated by mineral-associated SOM (> 95%). This SOM fraction comprised greater proportions of aryl- and carbonyl-C and is considered to contain a higher amount of microbially-derived organic substances. Land use can alter both, structure and stability of SOM fractions. All applied chemical treatments induced considerable SOC losses (> 70–95% of mineral-associated SOM) in the investigated soils. The proportion of residual C after chemical fractionation was largest in the arable Ap and E horizons and increased with decreasing C content in the initial SOC after stepwise hydrolysis as well as after the oxidative treatments with H2O2 and Na2S2O8. This can be expected for a functional stable pool of SOM, because it is assumed that the more easily available part of SOC is consumed first if C inputs decrease. All chemical treatments led to a preferential loss of the younger, maize-derived SOC, but this was most pronounced after the treatments with Na2S2O8 and H2O2. After all chemical fractionations, the mean 14C ages of SOC were higher than in the mineral-associated SOM fraction for both study sites and increased in the order: NaOCl < NaOCl+HF ≤ stepwise hydrolysis << H2O2 ≈ Na2S2O8. The results suggest that all treatments were capable of isolating a more stable SOM fraction, but the treatments with H2O2 and Na2S2O8 were the most efficient ones. However, none of the chemical fractionation methods was able to fit the IOM pool calculated using the Rothamsted Carbon Model and isotope data. In the evaluation of thermal oxidation for obtaining stable C fractions, SOC losses increased with temperature from 24–48% (200°C) to 100% (500°C). In the Halle maize Ap horizon, losses of the young, maize-derived C were considerably higher than losses of the older C3-derived C, leading to an increase in the apparent C turnover time from 220 years in mineral-associated SOC to 1158 years after thermal oxidation at 300°C. Most likely, the preferential loss of maize-derived C in the Halle soil was caused by the presence of the high amounts of fossil C mentioned above, which make up a relatively large thermally stable C3-C pool in this soil. This agrees with lower overall SOC losses for the Halle Ap horizon compared to the Rotthalmünster Ap horizon. In the Rotthalmünster soil only slightly more maize-derived than C3-derived SOC was removed by thermal oxidation. Apparent C turnover times increased slightly from 58 years in mineral-associated SOC to 77 years after thermal oxidation at 300°C in the Rotthalmünster Ap and from 151 to 247 years in the Rotthalmünster E horizon. This led to the conclusion that thermal oxidation of SOM was not capable of isolating SOM fractions of considerably higher stability. The incubation experiment showed that macroaggregates develop rapidly after the addition of easily available plant residues. Within the first four weeks of incubation, the maximum aggregation was reached in all treatments without addition of fungicide. The formation of water-stable macroaggregates was related to the size of the microbial biomass pool and its activity. Furthermore, fungi were found to be crucial for the development of soil macroaggregates as the formation of water-stable macroaggregates was significantly delayed in the fungicide treated soils. The C concentration in the obtained aggregate fractions decreased with decreasing aggregate size class, which is in line with the aggregate hierarchy postulated by several authors for soils with SOM as the major binding agent. Macroaggregation involved incorporation of large amounts maize-derived organic matter, but macroaggregates did not play the most important role in the stabilization of maize-derived SOM, because of their relatively low amount (less than 10% of the soil mass). Furthermore, the maize-derived organic matter was quickly incorporated into all aggregate size classes. The microaggregate fraction stored the largest quantities of maize-derived C and N – up to 70% of the residual maize-C and -N were stored in this fraction.

Effects of tillage on processes of organic matter sequestration

To increase the organic matter (OM) content in the soil is one main goal in arable soil management. The adoption of tillage systems with reduced tillage depth and/or frequency (reduced tillage) or of no-tillage was found to increase the concentration of soil OM compared to conventional tillage (CT; ploughing to 20-30 cm). However, the underlying processes are not yet clear and are discussed contradictorily. So far, few investigations were conducted on tillage systems with a shallow tillage depth (minimum tillage = MT; maximum tillage depth of 10 cm). A better understanding of the interactions between MT implementation and changes in OM transformation in soils is essential in order to evaluate the possible contribution of MT to a sustainable management of arable soils. The objectives of the present thesis were (i) to compare OM concentrations, microbial biomass, water-stable aggregates, and particulate OM (POM) between CT and MT soils, (ii) to estimate the temporal variability of water-stable aggregate size classes occurring in the field and the dynamics of macroaggregate (>250 µm) formation and disruption under controlled conditions, (iii) to investigate whether a lower disruption or a higher formation rate accounts for a higher occurrence of macroaggregates under MT compared to CT, (iv) to determine which fraction is the major agent for storing the surplus of OM found under MT compared to CT, and (v) to observe the early OM transformation after residue incorporation in different tillage systems simulated. Two experimental sites (Garte-Süd and Hohes Feld) near Göttingen, Germany, were investigated. Soil type of both sites was a Haplic Luvisol. Since about 40 years, both sites receive MT by a rotary harrow (to 5-8 cm depth) and CT by a plough (to 25 cm depth). Surface soils (0-5 cm) and subsoils (10-20 cm) of two sampling dates (after fallow and directly after tillage) were investigated for concentrations of organic C (Corg) and total N (N), different water-stable aggregate size classes, different density fractions (for the sampling date after fallow only), microbial biomass, and for biochemically stabilized Corg and N (by acid hydrolysis; for the sampling date after tillage only). In addition, two laboratory incubations were performed under controlled conditions: Firstly, MT and CT soils were incubated (28 days at 22°C) as bulk soil and with destroyed macroaggregates in order to estimate the importance of macroaggregates for the physical protection of the very labile OM against mineralization. Secondly, in a microcosm experiment simulating MT and CT systems with soil <250 µm and with 15N and 13C labelled maize straw incorporated to different depths, the mineralization, the formation of new macroaggregates, and the partitioning of the recently added C and N were followed (28 days at 15°C). Forty years of MT regime led to higher concentrations of microbial biomass and of Corg and N compared to CT, especially in the surface soil. After fallow and directly after tillage, a higher proportion of water-stable macroaggregates rich in OM was found in the MT (36% and 66%, respectively) than in the CT (19% and 47%, respectively) surface soils of both sites (data shown are of the site Garte-Süd only). The subsoils followed the same trend. For the sampling date after fallow, no differences in the POM fractions were found but there was more OM associated to the mineral fraction detected in the MT soils. A large temporal variability was observed for the abundance of macroaggregates. In the field and in the microcosm simulations, macroaggregates were found to have a higher formation rate after the incorporation of residues under MT than under CT. Thus, the lower occurrence of macroaggregates in CT soils cannot be attributed to a higher disruption but to a lower formation rate. A higher rate of macroaggregate formation in MT soils may be due to (i) the higher concentrated input of residues in the surface soil and/or (ii) a higher abundance of fungal biomass in contrast to CT soils. Overall, as a location of storage of the surplus of OM detected under MT compared to CT, water-stable macroaggregates were found to play a key role. In the incubation experiment, macroaggregates were not found to protect the very labile OM against mineralization. Anyway, the surplus of OM detected after tillage in the MT soil was biochemically degradable. MT simulations in the microcosm experiment showed a lower specific respiration and a less efficient translocation of recently added residues than the CT simulations. Differences in the early processes of OM translocation between CT and MT simulations were attributed to a higher residue to soil ratio and to a higher proportion of fungal biomass in the MT simulations. Overall, MT was found to have several beneficial effects on the soil structure and on the storage of OM, especially in the surface soil. Furthermore, it was concluded that the high concentration of residues in the surface soil of MT may alter the processes of storage and decomposition of OM. In further investigations, especially analysis of the residue-soil-interface and of effects of the depth of residue incorporation should be emphasised. Moreover, further evidence is needed on differences in the microbial community between CT and MT soils.

Bedeutung der C- und N-Rhizodeposition für das mikrobielle Wachstum und den mikrobiellen Umsatz von C und N in der Rhizosphäre in Abhängigkeit von der Entfernung zur Wurzel

Ziel dieser Arbeit war es, die Bedeutung des C und N mit Herkunft aus der Rhizodeposition für das mikrobielle Wachstum und den C- und N-Umsatz in der Rhizosphäre in Abhängigkeit von der Entfernung zur Wurzel zu untersuchen. Dazu wurde als wesentliche methodische Voraussetzung ein künstliches Rhizosphärensystem entwickelt, um die durch die Rhizodeposite induzierten Prozesse an der Grenzfläche zwischen Wurzeloberfläche und Boden zu untersuchen. Dieses eingesetzte Rhizosphärensystem wurde nach einem Vorbild eines in schon vielfältigen Untersuchungen der Rhizosphäre eingesetzten Systems von GAHOONIA und NIELSEN (1991) konzipiert. Zur Verfolgung der pflanzlichen C- und N-Rhizodeposition im Boden wurden 13C- und 15N-Tracer-Isotopen-Techniken zur Isotopenmarkierung der Testpflanzen eingesetzt. Zur Probenahme des Rhizosphärenbodens in räumlichen Abstand zur künstlichen Rhizoplane wurde eine Schneidvorrichtung nach FITZ et al. (2003) eingesetzt, die es ermöglicht frische Bodenproben in definiertem Abstand zur künstlichen Rhizoplane zu schneiden. Das Rhizosphärensystem wurde in 13C- und 15N-Doppelmarkierungsexperimenten mit Lolium perenne, Triticum aestivum und Avena sativa eingesetzt.Das unterschiedliche Ansprechen der mikrobiellen Gemeinschaft auf den Substrateintrag in unterschiedlicher Entfernung zur Wurzel zeigte komplexe Wechselwirkungen in Bezug auf das mikrobielle Wachstum, den mikrobiellen Umsatz, den Substrateintrag aus der Rhizodeposition und den stimulierten Abbau der nativen organischen Bodensubstanz. Der Eintrag von Rhizodepositen stellt daher eine bedeutende Funktion in der Regulation der mikrobiellen Biomasse und der Prozesse des Umsatzes der organischen Bodensubstanz während und wahrscheinlich bis nach der Vegetationsperiode dar. Die simultane Verfolgung von Rhizodepositions-C und -N im Boden und deren Funktionen innerhalb der Rhizosphäre, gerade im Hinblick auf die mikrobielle Biomasse und den Umsatz der organischen Substanz im Boden, führt zu einem besseren Verständnis der komplexen wechselseitigen Prozesse zwischen Pflanze und Boden.

Leguminous cover crops differentially affect maize yields in three contrasting soil types of Kakamega, Western Kenya

Maize production in smallholder farming systems in Kenya is largely limited by low soil fertility. As mineral fertilizer is expensive, green manuring using leguminous cover crops could be an alternative strategy for farmers to enhance farm productivity. However due to variability in soil type and crop management, the effects of green manure are likely to differ with farms. The objectives of this study were to evaluate Mucuna pruriens and Arachis pintoi on (i) biomass and nitrogen fixation (^15N natural abundance), (ii) soil carbon and nitrogen stocks and (iii) their effects on maize yields over two cropping seasons in Kakamega, Western Kenya. Mucuna at 6 weeks accumulated 1–1.3 Mg ha^{-1} of dry matter and 33–56 kg ha^{-1} nitrogen of which 70% was nitrogen derived from the atmosphere (Ndfa). Arachis after 12 months accumulated 2–2.7 Mg ha^{-1} of dry matter and 51–74 kg N ha^{-1} of which 52-63 % was from Ndfa. Soil carbon and nitrogen stocks at 0–15 cm depth were enhanced by 2-4 Mg C ha^{-1} and 0.3–1.0 Mg N ha^{-1} under Mucuna and Arachis fallow, irrespective of soil type. Maize yield increased by 0.5-2 Mg ha^{-1} in Mucuna and 0.5–3 Mg ha^{-1} in Arachis and the response was stronger on Nitisol than on Acrisol or Ferralsol. We concluded that leguminous cover crops seem promising in enhancing soil fertility and maize yields in Kenya, provided soil conditions and rainfall are suitable.

Multi-isotopic and compositional exploration of factors controlling nitrate pollution

In Catalonia, according to the nitrate directive (91/676/EU), nine areas have been declared as vulnerable to nitrate pollution from agricultural sources (Decret 283/1998 and Decret 479/2004). Five of these areas have been studied coupling hydro chemical data with a multi-isotopic approach (Vitòria et al. 2005, Otero et al. 2007, Puig et al. 2007), in an ongoing research project looking for an integrated application of classical hydrochemistry data, with a comprehensive isotopic characterisation (δ15N and δ18O of dissolved nitrate, δ34S and δ18O of dissolved sulphate, δ13C of dissolved inorganic carbon, and δD and δ18O of water). Within this general frame, the contribution presented explores compositional ways of: (i) distinguish agrochemicals and manure N pollution, (ii) quantify natural attenuation of nitrate (denitrification), and identify possible controlling factors. To achieve this two-fold goal, the following techniques have been used. Separate biplots of each suite of data show that each studied region has a distinct δ34S and pH signatures, but they are homogeneous with regard to NO3- related variables. Also, the geochemical variables were projected onto the compositional directions associated with the possible denitrification reactions in each region. The resulting balances can be plot together with some isotopes, to assess their likelihood of occurrence

Stable Isotope Analysis Reveals That Agricultural Habitat Provides an Important Dietary Component for Nonbreeding Dunlin

Although shorebirds spending the winter in temperate areas frequently use estuarine and supratidal (upland) feeding habitats, the relative contribution of each habitat to individual diets has not been directly quantified. We quantified the proportional use that Calidris alpina pacifica (Dunlin) made of estuarine vs. terrestrial farmland resources on the Fraser River Delta, British Columbia, using stable isotope analysis (δ13C, δ15N) of blood from 268 Dunlin over four winters, 1997 through 2000. We tested for individual, age, sex, morphological, seasonal, and weather-related differences in dietary sources. Based on single- (δ13C) and dual-isotope mixing models, the agricultural habitat contributed approximately 38% of Dunlin diet averaged over four winters, with the balance from intertidal flats. However, there was a wide variation among individuals in the extent of agricultural feeding, ranging from about 1% to 95% of diet. Younger birds had a significantly higher terrestrial contribution to diet (43%) than did adults (35%). We estimated that 6% of adults and 13% of juveniles were obtaining at least 75% of their diet from terrestrial sources. The isotope data provided no evidence for sex or overall body size effects on the proportion of diet that is terrestrial in origin. The use of agricultural habitat by Dunlin peaked in early January. Adult Dunlin obtained a greater proportion of their diet terrestrially during periods of lower temperatures and high precipitation, whereas no such relationship existed for juveniles. Seasonal variation in the use of agricultural habitat suggests that it is used more during energetically stressful periods. The terrestrial farmland zone appears to be consistently important as a habitat for juveniles, but for adults it may provide an alternative feeding site used as a buffer against starvation during periods of extreme weather. Loss or reduction of agricultural habitat adjacent to estuaries may negatively impact shorebird fitness, with juveniles disproportionately affected.

Establishing Winter Origins of Migrating Lesser Snow Geese Using Stable Isotopes

Increases in Snow Goose (Chen caerulescens) populations and large-scale habitat changes in North America have contributed to the concentration of migratory waterfowl on fewer wetlands, reducing resource availability, and enhancing risks of disease transmission. Predicting wintering locations of migratory individuals is critical to guide wildlife population management and habitat restoration. We used stable carbon (δ13C), nitrogen (δ15N), and hydrogen (δ2H) isotope ratios in muscle tissue of wintering Snow Geese to discriminate four major wintering areas, the Playa Lake Region, Texas Gulf Coast, Louisiana Gulf Coast, and Arkansas, and infer the wintering locations of individuals collected later during the 2007 and 2008 spring migrations in the Rainwater Basin (RWB) of Nebraska. We predicted the wintering ground derivation of migrating Snow Geese using a likelihood-based approach. Our three-isotope analysis provided an efficient discrimination of the four wintering areas. The assignment model predicted that 53% [95% CI: 37-69] of our sample of Snow Geese from the RWB in 2007 had most likely originated in Louisiana, 38% [23-54] had wintered on Texas Gulf Coast, and 9% [0-20] in Arkansas; the assessment suggested that 89% [73-100] of our 2008 sample had most likely come from Texas Gulf Coast, 9% [0-27] from Louisiana Gulf Coast, and 2% [0-9] from Arkansas. Further segregation of wintering grounds and additional sampling of spring migrating Snow Geese would refine overall assignment and help explain interannual variations in migratory connectivity. The ability to distinguish origins of northbound geese can support the development of spatially-adaptive management strategies for the midcontinent Snow Goose population. Establishing migratory connectivity using isotope assignment techniques can be extended to other waterfowl species to determine critical habitat, evaluate population energy requirements, and inform waterfowl conservation and management strategies.

West African storm tracks and their relationship to Atlantic tropical cyclones

The automatic tracking technique used by Thorncroft and Hodges (2001) has been used to identify coherent vorticity structures at 850hPa over West Africa and the tropical Atlantic in the ECMWF 40-year reanalysis. The presence of two dominant source regions, north and south of 15ºN over West Africa, for storm tracks over the Atlantic was confirmed. Results show that the southern storm track provides most of the storms that reach the main development region where most tropical cyclones develop. There exists marked seasonal variability in location and intensity of the storms leaving the West African coast, which may influence the likelihood of downstream intensification and longevity. There exists considerable year-to-year variability in the number of West African storm tracks, both in numbers over the land and continuing out over the tropical Atlantic Ocean. While the low-frequency variability is well correlated with Atlantic tropical cyclone activity, West African rainfall and SSTs, the interannual variability is found to be uncorrelated with these. In contrast, variance of the 2-6-day-filtered meridional wind, which provides a synoptic-scale measure of African Easterly Wave activity, shows a significant, positive correlation with tropical cyclone activity at interannual timescales.