Soil property and management effects on grassland microbial communities across a latitudinal gradient in Germany

There is much interest in the identification of the main drivers controlling changes in the microbial community that may be related to sustainable land use. We examined the influence of soil properties and land-use intensity (N fertilization, mowing, grazing) on total phospholipid fatty acid (PLFA) biomass, microbial community composition (PLFA profiles) and activities of enzymes involved in the C, N, and P cycle. These relationships were examined in the topsoil of grasslands from three German regions (Schorfheide-Chorin (SCH), Hainich-Dun (HAI), Schwabische Alb (ALB)) with different parent material. Differences in soil properties explained 60% of variation in PLFA data and 81% of variation in enzyme activities across regions and land-use intensities. Degraded peat soils in the lowland areas of the SCH with high organic carbon (OC) concentrations and sand content contained lower PLFA biomass, lower concentrations of bacterial, fungal, and arbuscular mycorrhizal PLFAs, but greater enzyme activities, and specific enzyme activities (per unit microbial biomass) than mineral soils in the upland areas of the HAI and ALB, which are finer textured, drier, and have smaller OC concentrations. After extraction of variation that originated from large-scale differences among regions and differences in land-use intensities between plots, soil properties still explained a significant amount of variation in PLFA data (34%) and enzyme activities (60%). Total PLFA biomass and all enzyme activities were mainly related to OC concentration, while relative abundance of fungi and fungal to bacterial ratio were mainly related to soil moisture. Land-use intensity (LUI) significantly decreased the soil C:N ratio. There was no direct effect of LUI on total PLFA biomass, microbial community composition, N and P cycling enzyme activities independent of study region and soil properties. In contrast, the activities and specific activities of enzymes involved in the C cycle increased significantly with LUI independent of study region and soil properties, which can have impact on soil organic matter decomposition and nutrient cycling. Our findings demonstrate that microbial biomass and community composition as well as enzyme activities are more controlled by soil properties than by grassland management at the regional scale. (C) 2013 Elsevier B.V: All rights reserved.

Biotic and Abiotic Properties Mediating Plant Diversity Effects on Soil Microbial Communities in an Experimental Grassland

Plant diversity drives changes in the soil microbial community which may result in alterations in ecosystem functions. However, the governing factors between the composition of soil microbial communities and plant diversity are not well understood. We investigated the impact of plant diversity (plant species richness and functional group richness) and plant functional group identity on soil microbial biomass and soil microbial community structure in experimental grassland ecosystems. Total microbial biomass and community structure were determined by phospholipid fatty acid (PLFA) analysis. The diversity gradient covered 1, 2, 4, 8, 16 and 60 plant species and 1, 2, 3 and 4 plant functional groups (grasses, legumes, small herbs and tall herbs). In May 2007, soil samples were taken from experimental plots and from nearby fields and meadows. Beside soil texture, plant species richness was the main driver of soil microbial biomass. Structural equation modeling revealed that the positive plant diversity effect was mainly mediated by higher leaf area index resulting in higher soil moisture in the top soil layer. The fungal-to-bacterial biomass ratio was positively affected by plant functional group richness and negatively by the presence of legumes. Bacteria were more closely related to abiotic differences caused by plant diversity, while fungi were more affected by plant-derived organic matter inputs. We found diverse plant communities promoted faster transition of soil microbial communities typical for arable land towards grassland communities. Although some mechanisms underlying the plant diversity effect on soil microorganisms could be identified, future studies have to determine plant traits shaping soil microbial community structure. We suspect differences in root traits among different plant communities, such as root turnover rates and chemical composition of root exudates, to structure soil microbial communities.

Evaluación de las características físico-químicas y biológicas en dos suelos superficiales cultivados con pera (Pyrus communis L.) cv. Williams bajo manejo convencional

El objetivo del trabajo fue identificar las características físico-químicas y biológicas en dos suelos superficiales fertilizados con nitrógeno y enmienda orgánica en el Alto Valle de Río Negro (huertos H1 y H2). En ambos huertos se aplicó fertilizante nitrogenado durante las temporadas 2008-2009 y 2009-2010 y en H2 se aplicó estiércol de pollo. Se extrajeron muestras de suelos en primavera y otoño y se determinó: carbono orgánico total, conductividad eléctrica, cationes de intercambio, relación de adsorción de sodio y nitratos, respiración microbiana, carbono de la biomasa microbiana, actividad de la deshidrogenasa y el índice de mineralización. La concentración de carbono orgánico total, potasio y nitrógeno fueron adecuadas para la producción de pera. El comportamiento de las variables biológicas fue diferente en los huertos. En H1 fueron mayores en primavera, en ambas temporadas y el índice de mineralización fue ligeramente superior a 1 en otoño, indicando equilibrio entre la mineralización y la humificación del carbono. En H2 las mediciones biológicas fueron similares entre las estaciones como consecuencia de realizar fertilización nitrogenada (N) en primavera y en otoño. La enmienda orgánica no reflejó un aumento de la actividad biológica en primavera. La actividad microbiana y enzimática en H1 y H2 fue sensible a los cambios que ocurrieron en los suelos.

Abundance, biomass, production, grazing, and biometric measurements of prokaryotes, nanoflagellates, ciliates, and dinoflagellates in three areas close to the Antarctic Peninsula in spring and summer 1995/96

Two cruises were carried out during the Austral spring-summer (November 1995 - January 1996: FRUELA 95, and January - February 1996: FRUELA 96), sampling in Bellingshausen Sea, western Bransfield Strait and Gerlache Strait. We investigated whether there were any spatial (among locations) or temporal (between cruises) differences in abundance and biomass of microbial heterotrophic and autotrophic assemblages. Changes in the concentration of chlorophyll a, prokaryotes, heterotrophic and phototrophic nanoflagellates abundance and biomass were followed in the above mentioned locations close to the Antarctic Peninsula. Parallel to these measurements we selected seven stations to determine grazing rates on prokaryotes by protists at a depth coincident with the depth of maximum chlorophyll a concentration. Measuring the disappearance of fluorescent minicells over 48 h assessed grazing by the protist community. From prokaryotes grazing rates, we estimated how much prokaryotic carbon was channeled to higher trophic levels (protists), and whether this prokaryotic carbon could maintain protists biomass and growth rates. In general higher values were reported for Gerlache Strait than for the other two areas. Differences between cruises were more evident for the oligotrophic areas in Bellingshausen Sea and Bransfield Strait than in Gerlache Strait (eutrophic area). Higher values for phototrophic (at least for chlorophyll a concentration) and abundance of all heterotrophic microbial populations were recorded in Bellingshausen Sea and Bransfield Strait during late spring - early summer (FRUELA 95) than in mid-summer (FRUELA 96). However, similar results for these variables were observed in Gerlache Strait as in spring-early summer as well as in mid-summer. Also, we found differences in grazing rates on prokaryotes among stations located in the three areas and between cruises. Thus, during late spring-early summer (FRUELA 95), the prokaryotic biomass consumed from the standing stock was higher in Bellingshausen Sea (26%/day) and Gerlache Strait (18-26%/day) than in Bransfield Strait (0.68-14%/day). During mid-summer (FRUELA 96) a different pattern was observed. The station located in Bellingshausen Sea showed higher values of prokaryotic biomass consumed (11%/day) than the one located in Gerlache Strait (2.3%/day). Assuming HNF as the main prokaryotic consumers, we estimated that the prokaryotic carbon consumed by heterotrophic nanoflagellates (HNF) barely covers their carbon requirements for growth. These results suggest that in Antarctic waters, HNF should feed in other carbon sources than prokaryotes.

Plankton biomass, POC and PON concentrations, and sinking velocities during the 2008 spring diatom bloom in Disko Bay

Phytoplankton and copepod succession was investigated in Disko Bay, western Greenland from February to July 2008. The spring phytoplankton bloom developed immediately after the breakup of sea ice and reached a peak concentration of 24 mg chl a/m**3 2 wk later. The bloom was analyzed during 3 phases: the developing, the decaying, and the post-bloom phases. Grazing impact by the copepod community was assessed by 4 methods; gut fluorescence, in situ faecal pellet production, and egg and faecal pellet production from bottle incubations. Calanus spp. dominated the mesozooplankton community. They were present from the initiation of the bloom but only had a small grazing impact on the phytoplankton. Consequently, there was a close coupling between the spring phytoplankton bloom and sedimentation of particulate organic carbon (POC). Out of 1836 ±180 mg C/m**2/d leaving the upper 50 m, 60 % was phytoplankton based carbon (PPC). The composition and quality of the sedimenting material changed throughout the bloom succession from PPC dominance in the initial phase with a POC/PON ratio close to 6.6 to a dominance of amorphous detritus with a higher POC/PON ratio (>10) in the post-bloom phase. The succession and fate of the phytoplankton spring bloom was controlled by nitrogen limitation and subsequent sedimentation, while grazing-mediated flux by the Calanus-dominated copepod community played a minor role in the termination of the spring bloom of Disko Bay.

Zooplankton data of M83/1

Oxygen-deficient waters in the ocean, generally referred to as oxygen minimum zones (OMZ), are expected to expand as a consequence of global climate change. Poor oxygenation is promoting microbial loss of inorganic nitrogen (N) and increasing release of sediment-bound phosphate (P) into the water column. These intermediate water masses, nutrient-loaded but with an N deficit relative to the canonical N:P Redfield ratio of 16:1, are transported via coastal upwelling into the euphotic zone. To test the impact of nutrient supply and nutrient stoichiometry on production, partitioning and elemental composition of dissolved (DOC, DON, DOP) and particulate (POC, PON, POP) organic matter, three nutrient enrichment experiments were conducted with natural microbial communities in shipboard mesocosms, during research cruises in the tropical waters of the southeast Pacific and the northeast Atlantic. Maximum accumulation of POC and PON was observed under high N supply conditions, indicating that primary production was controlled by N availability. The stoichiometry of microbial biomass was unaffected by nutrient N:P supply during exponential growth under nutrient saturation, while it was highly variable under conditions of nutrient limitation and closely correlated to the N:P supply ratio, although PON:POP of accumulated biomass generally exceeded the supply ratio. Microbial N:P composition was constrained by a general lower limit of 5:1. Channelling of assimilated P into DOP appears to be the mechanism responsible for the consistent offset of cellular stoichiometry relative to inorganic nutrient supply and nutrient drawdown, as DOP build-up was observed to intensify under decreasing N:P supply. Low nutrient N:P conditions in coastal upwelling areas overlying O2-deficient waters seem to represent a net source for DOP, which may stimulate growth of diazotrophic phytoplankton. These results demonstrate that microbial nutrient assimilation and partitioning of organic matter between the particulate and the dissolved phase are controlled by the N:P ratio of upwelled nutrients, implying substantial consequences for nutrient cycling and organic matter pools in the course of decreasing nutrient N:P stoichiometry.

Biogeochemistry of mesocosm experiments in the tropical Pacific and Atlantic Ocean

Oxygen-deficient waters in the ocean, generally referred to as oxygen minimum zones (OMZ), are expected to expand as a consequence of global climate change. Poor oxygenation is promoting microbial loss of inorganic nitrogen (N) and increasing release of sediment-bound phosphate (P) into the water column. These intermediate water masses, nutrient-loaded but with an N deficit relative to the canonical N:P Redfield ratio of 16:1, are transported via coastal upwelling into the euphotic zone. To test the impact of nutrient supply and nutrient stoichiometry on production, partitioning and elemental composition of dissolved (DOC, DON, DOP) and particulate (POC, PON, POP) organic matter, three nutrient enrichment experiments were conducted with natural microbial communities in shipboard mesocosms, during research cruises in the tropical waters of the southeast Pacific and the northeast Atlantic. Maximum accumulation of POC and PON was observed under high N supply conditions, indicating that primary production was controlled by N availability. The stoichiometry of microbial biomass was unaffected by nutrient N:P supply during exponential growth under nutrient saturation, while it was highly variable under conditions of nutrient limitation and closely correlated to the N:P supply ratio, although PON:POP of accumulated biomass generally exceeded the supply ratio. Microbial N:P composition was constrained by a general lower limit of 5:1. Channelling of assimilated P into DOP appears to be the mechanism responsible for the consistent offset of cellular stoichiometry relative to inorganic nutrient supply and nutrient drawdown, as DOP build-up was observed to intensify under decreasing N:P supply. Low nutrient N:P conditions in coastal upwelling areas overlying O2-deficient waters seem to represent a net source for DOP, which may stimulate growth of diazotrophic phytoplankton. These results demonstrate that microbial nutrient assimilation and partitioning of organic matter between the particulate and the dissolved phase are controlled by the N:P ratio of upwelled nutrients, implying substantial consequences for nutrient cycling and organic matter pools in the course of decreasing nutrient N:P stoichiometry.

Analysis of spatial microbial properties from experimental plots of the Jena experiment (September 2010)

The study was carried out on the main plots (Main Experiment) of a large grassland biodiversity experiment, the Jena Experiment. In the main experiment, 82 grassland plots of 20 x 20 m were established from a pool of 60 species belonging to four functional groups (grasses, legumes, tall and small herbs). In May 2002, varying numbers of plant species from this species pool were sown into the plots to create a gradient of plant species richness (1, 2, 4, 8, 16 and 60 species) and functional richness (1, 2, 3, 4 functional groups). Plots were maintained by bi-annual weeding and mowing. This data set consists of standard deviation (SD), mean and stability (stab) of soil microbial basal respiration (µl O2/h/g dry soil) and microbial biomass carbon (µg C/g dry soil). Data were derived by taking soil samples and measuring basal and substrate-induced microbial respiration with an oxygen-consumption apparatus. Samples for calculating the spatial stability of soil microbial properties were taken on the 20th of September in 2010. Oxygen consumption of soil microorganisms in fresh soil equivalent to 3.5 g dry weight was measured at 22°C over a period of 24 h. Basal respiration (µlO2/g dry soil/h) was calculated as mean of the oxygen consumption rates of hours 14 to 24 after the start of measurements. Substrate- induced respiration was determined by adding D-glucose to saturate catabolic enzymes of microorganisms according to preliminary studies (4 mg g-1 dry soil solved in 400 µl deionized water). Maximum initial respiratory response (µl O2/g dry soil/ h) was calculated as mean of the lowest three oxygen consumption values within the first 10 h after glucose addition. Microbial biomass carbon (µg C/g dry soil) was calculated as 38 × Maximum initial respiratory response according to prelimiray studies.

Analysis of temporal microbial properties from experimental plots of the Jena experiment (2003-2014)

The study was carried out on the main plots (Main Experiment) of a large grassland biodiversity experiment, the Jena Experiment. In the main experiment, 82 grassland plots of 20 x 20 m were established from a pool of 60 species belonging to four functional groups (grasses, legumes, tall and small herbs). In May 2002, varying numbers of plant species from this species pool were sown into the plots to create a gradient of plant species richness (1, 2, 4, 8, 16 and 60 species) and functional richness (1, 2, 3, 4 functional groups). Plots were maintained by bi-annual weeding and mowing. This data set consists of standard deviation (SD), mean and stability (stab) of soil microbial basal respiration (µl O2/h/g dry soil) and microbial biomass carbon (µg C/g dry soil). Data were derived by taking soil samples and measuring basal and substrate-induced microbial respiration with an oxygen-consumption apparatus. Samples for calculating the temporal stability were taken every year in May/June from 2003 to 2014, except in 2005. Oxygen consumption of soil microorganisms in fresh soil equivalent to 3.5 g dry weight was measured at 22°C over a period of 24 h. Basal respiration (µlO2/g dry soil/h) was calculated as mean of the oxygen consumption rates of hours 14 to 24 after the start of measurements. Substrate- induced respiration was determined by adding D-glucose to saturate catabolic enzymes of microorganisms according to preliminary studies (4 mg g-1 dry soil solved in 400 µl deionized water). Maximum initial respiratory response (µl O2/g dry soil/h) was calculated as mean of the lowest three oxygen consumption values within the first 10 h after glucose addition. Microbial biomass carbon (µg C/g dry soil) was calculated as 38 × Maximum initial respiratory response according to prelimiray studies.

Flujos de gases de efecto invernadero en suelos forestales y agroforestales del centro de la Península Ibérica=Greenhouse gas flows in forest and agroforestry soils in the center of the Iberian Peninsula

Debido a la complejidad de los procesos que controlan el intercambio de gases de carbono (C) y nitrógeno (N) entre el suelo y la atmósfera, en los sistemas forestales y agroforestales, son comprensibles las incógnitas existentes respecto a la estimación de los flujos de los gases de efecto invernadero (GEI) y la capacidad como reservorios de carbono de los suelos, bajo diferentes formas de uso y regímenes de alteración a escala regional y global. Esta escasez de información justifica la necesidad de caracterizar la dinámica de intercambio de GEI en los ecosistemas Mediterráneos, en especial en el contexto actual de cambio climático, y el incremento asociado de temperatura y periodos de sequía, alteración de los patrones de precipitación, y el riesgo de incendios forestales; cuyas consecuencias afectarán tanto a los compartimentos de C y de N del suelo como a la capacidad de secuestro de C de estos ecosistemas. Dentro de este contexto se enmarca la presente tesis doctoral cuyo objetivo ha sido cuantificar y caracterizar los flujos de dióxido de carbono (CO2), de oxido nitroso (N2O) y de metano (CH4), junto con los stocks de C y N, en suelos forestales de Quercus ilex, Quercus pyrenaica y Pinus sylvestris afectados por incendios forestales; así como el estudiar el efecto de la gestión y la cubierta arbórea en la respiración del suelo y los stocks de C y N en una dehesa situada en el centro de la Península Ibérica. De manera que los flujos de CO2, N2O y CH4; y los parámetros físico-químicos y biológicos del suelo fueron estudiados en los diferentes tratamientos y ecosistemas a lo largo del trabajo que se presenta. Los resultados obtenidos muestran la existencia de variaciones temporales y espaciales de la respiración del suelo dentro de una escala geográfica pequeña, controladas principalmente por la temperatura y la humedad del suelo; y por los contenidos de C y N del suelo en un bosque de Pinus sylvestris en la vertiente norte de la Sierra de Guadarrama , en España. El análisis de los efectos de los incendios forestales a largo plazo (6-8 años) revela que las pérdidas anuales de C a través de la respiración del suelo en las zonas quemadas de Quercus ilex, Quercus pyrenaica y Pinus sylvestris fueron 450 gCm-2yr-1, 790 gCm-2yr-1 y 1220 gCm-2yr-1, respectivamente; lo que representa una reducción del 43%, 22% y 11% en comparación con las zonas no quemadas de dichas especies, debido a la destrucción de la masa arbórea. El efecto del fuego también alteró los flujos N2O y CH4 del suelo, de una forma diferente en los distintos ecosistemas y estacionalidades estudiadas. De tal modo, que los suelos quemados mostraron una mayor oxidación del CH4 en las masas de Q. ilex, y una menor oxidación en las de P. sylvestris; además de una disminución de los flujos de N2O en Q. pyrenaica. Los incendios también afectaron los parámetros microclimáticos de los suelos forestales, observándose un incremento de la temperatura del suelo y una disminución de la humedad en los emplazamientos quemados que en los no quemados. Los cationes intercambiables, el pH, el cociente C/N, el contenido en raicillas y la biomasa microbiana también disminuyeron en las zonas quemadas. Aunque el C orgánico del suelo no se alteró de manera significativa, si lo hizo la calidad de la materia orgánica, disminuyendo el carbono lábil y aumentando las formas recalcitrantes lo que se tradujo en menor sensibilidad de la respiración del suelo a la temperatura (valores de Q10) en las zonas quemadas. Los resultados del estudio realizado en la Dehesa muestran que las actividades silvopastorales estudiadas afectaron levemente y de forma no constante a la respiración del suelo y las condiciones microclimáticas del suelo. Se observó una reducción 12% de la respiración del suelo por efecto del pastoreo no intensivo. Sin embargo, se observaron incrementos de 3Mg/ha en los stocks de C y de 0.3 Mg/ha en los stocks de N en los suelos pastoreados en comparación con los no pastoreados. Aunque, no se observó un claro efecto de la labranza sobre la respiración del suelo en nuestro experimento, sin embargo si se observó una disminución de 3.5 Mg/ha en las reservas de C y de 0.3 Mg/ ha en las de N en los suelos labrados comparados con los no labrados. La copa del arbolado influyó de forma positiva tanto en la respiración del suelo, como en los stocks de C y N de los suelos. La humedad del suelo jugó un papel relevante en la sensibilidad de la respiración a la temperatura del suelo. Nuestros resultados ponen de manifiesto la sensibilidad de la respiración del suelo a cambios en la humedad y los parámetros edáficos, y sugieren que la aplicación de modelos estándar para estimar la respiración del suelo en áreas geográficas pequeñas puede no ser adecuada a menos que otros factores sean considerados en combinación con la temperatura del suelo. Además, las diferentes respuestas de los flujos de gases de efecto invernadero a los cambios, años después de la ocurrencia de incendios forestales, destaca la necesidad de incluir estos cambios en las futuras investigaciones de la dinámica del carbono en los ecosistemas mediterráneos. Por otra parte, las respuestas divergentes en los valores de respiración del suelo y en los contenidos de C y N del suelo observados en la dehesa, además de la contribución de la copa de los árboles en los nutrientes del suelo ilustran la importancia de mantener la gestión tradicional aplicada en beneficio de la capacidad de almacenar C en la dehesa estudiada. La información obtenida en este trabajo pretende contribuir a la mejora del conocimiento de la dinámica y el balance de C en los sistemas mediterráneos, además de ayudar a predecir el impacto del cambio climático en el intercambio de C entre los ecosistemas forestales y agroforestales y la atmósfera. ABSTRACT Due to the complexity of the processes that control the exchange of carbon (C) and nitrogen (N) gasses between soils and the atmosphere in forest and agroforestry ecosystems, understandable uncertainties exist as regards the estimation of greenhouse gas (GHG) fluxes and the soil sink capacity at regional and global scale under different forms of land use and disturbance regimes. These uncertainties justify the need to characterize the exchange dynamics of GHG between the atmosphere and soils in Mediterranean terrestrial ecosystems, particularly in the current context of climate change and the associated increase in temperature, drought periods, heavy rainfall events, and increased risk of wildfires, which affect not only the C and N pools but also the soil C sink capacity of these ecosystems. Within this context, the aims of the present thesis were, firstly, to quantify and characterize the fluxes of carbon dioxide (CO2), nitrous oxide (N2O) and methane (CH4) as well as the C and N stocks in Quercus ilex, Quercus pyrenaica and Pinus sylvestris stands affected by wildfires, and secondly, to study the effects of Quercus ilex canopy and management on both soil respiration and C and N pools in dehesa systems in the center of Iberian Peninsula. Soil CO2, N2O and CH4 fluxes, and soil physical-chemical and biological parameters were studied under the different treatments and ecosystems considered in this study. The results showed seasonal and spatial variations in soil respiration within small geographic areas, mainly controlled by soil temperature and moisture in addition to soil carbon and nitrogen stocks in mixed pine–oak forest ecosystems on the north facing slopes of the Sierra de Guadarrama in Spain. The analysis of long term effects of wildfires (6–8 years) revealed that annual carbon losses through soil respiration from burned sites in Quercus ilex, Quercus pyrenaica and Pinus sylvestris stands were 450 gCm-2yr-1, 790 gCm-2yr-1 and 1220 gCm-2yr-1, respectively; with burned sites emitting 43%, 22% and 11% less in burned as opposed to non-burned sites due the loss of trees. Fire may alter both N2O and CH4 fluxes although the magnitude of such variation depends on the site, soil characteristics and seasonal climatic conditions. The burned sites showed higher CH4 oxidation in Q.ilex stands, and lower oxidation rates in P. sylvestris stands. A reduction in N2O fluxes in Q. pyrenaica stands was detected at burned sites along with changes in soil microclimate; higher soil temperature and lower soil moisture content. Exchangeable cations, the C/N ratio, pH, fine root and microbial biomass were also found to decrease at burned sites. Although the soil organic carbon was not significantly altered, the quality of the organic matter changed, displaying a decrease in labile carbon and a relative increase in refractory forms, leading to lower sensitivity of soil respiration to temperature (Q10 values) at burned sites. The results from the dehesa study show that light grazing and superficial tilling practices used in the studied dehesa system in Spain had a slight but non-consistent impact on soil respiration and soil microclimate over the study period. The reduction in soil respiration in the dehesa system due to the effects of grazing was around 12 %. However, increments of 3Mg/ha in C stocks and 0.3 Mg/ha in N stocks in grazed soils were observed. Although no clear effect of tilling on soil respiration was found, a decrease of 3.5 Mg/ha in C stocks and 0.3 Mg/ha in N stocks was detected for tilled soils. The presence of a tree canopy induced increases in soil respiration, soil C and N stocks, while soil moisture was found to play an important role in soil respiration temperature response. Our results suggest that the use of standard models to estimate soil respiration in small geographical areas may not be adequate unless other factors are considered in addition to soil temperature. Furthermore, the different responses of GHG flux to climatic shifts, many years after the occurrence of wildfire, highlight the need to include these shifts in C dynamics in future research undertaken in Mediterranean ecosystems. Furthermore, divergent responses in soil respiration and soil C and N stocks to grazing or tilling practices in Dehesa systems, and the influence of tree canopy on soil respiration and soil nutrient content, illustrate the importance of maintaining beneficial management practices. Moreover, the carbon sequestration capacity of the Dehesa system studied may be enhanced through improvements in the management applied. It is hoped that the information obtained through this research will contribute towards improving our understanding of the dynamics and balance of C in Mediterranean systems, and help predict the impact of climate change on the exchange of C between forest and agroforestry ecosystems and the atmosphere.

Activity and biomass of small benthic biota in the central Arctic Ocean

Sediment samples collected during the expedition "Arctic Ocean '96" with the Swedish ice-breaker ODEN were investigated to estimate for the first time heterotrophic activity and total microbial biomass (size range from bacteria to small metazoans) from the perennially ice-covered central Arctic Ocean. Benthic activities and biomass were evaluated analysing a series of biogenic sediment compounds (i.e. bacterial exoenzymes, total adenylates, DNA, phospholipids, particulate proteins). In contrast to the very time-consuming sorting, enumeration and weight determination, analyses of biochemical sediment parameters may represent a useful method for obtaining rapid information on the ecological situation in a given benthic system. Bacterial cell numbers and biomass were estimated for comparison with biochemically determined biomass data, to evaluate the contribution of the bacterial biomass to the total microbial biomass. It appeared that bacterial biomass made up only 8-31% (average of all stations = 20%) of the total microbial biomass, suggesting a large fraction of other small infaunal organisms within the sediment samples (most probably fungi, yeasts, protozoans such as flagellates, ciliates or amoebae, as well as a fraction of small metazoans). Activity and biomass values determined within this study were generally extremely low, and often even slightly lower than those given for other deep oceanic regions, thus characterizing the seafloor of the central Arctic Ocean as a "benthic desert". Nevertheless, some clear trends in the data could be found, e.g. generally sharply decreasing values within the sediment column, a vague tendency for declining values with increasing water depth of sampling stations, and also differences between various Arctic deep-sea regions.